Hindawi Publishing Corporation e Scientific World Journal Volume 2014, Article ID 296590, 12 pages http://dx.doi.org/10.1155/2014/296590

Research Article Genetic Biodiversity of Italian Olives (Olea europaea) Germplasm Analyzed by SSR Markers Innocenzo Muzzalupo,1 Giuseppe Giovanni Vendramin,2 and Adriana Chiappetta3 1

Consiglio per la Ricerca e Sperimentazione per l’Agricoltura, Centro di Ricerca per l’Olivicoltura e l’Industria Olearia (CRA-OLI), C.da Li Rocchi-Vermicelli, 87036 Rende, Italy 2 Consiglio Nazionale delle Ricerche, Istituto di Bioscienze e BioRisorse, 50019 Sesto Fiorentino, Italy 3 Universit`a della Calabria, Dipartimento di Biologia, Ecologia e Scienza della Terra, Ponte P. Bucci, 87036 Arcavacata di Rende, Italy Correspondence should be addressed to Innocenzo Muzzalupo; [email protected] Received 19 December 2013; Accepted 22 January 2014; Published 27 February 2014 Academic Editors: M. Cresti, F. Grassi, and J. Jakse Copyright © 2014 Innocenzo Muzzalupo et al. This is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. The olive is an important fruit species cultivated for oil and table olives in Italy and the Mediterranean basin. The conservation of cultivated plants in ex situ collections is essential for the optimal management and use of their genetic resources. The largest ex situ olive germplasm collection consists of approximately 500 Italian olive varieties and corresponding to 85% of the total Italian olive germplasm is maintained at the Consiglio per la Ricerca e sperimentazione per l’Agricoltura, Centro di Ricerca per l’Olivicoltura e l’Industria Olearia (CRA-OLI), in Italy. In this work, eleven preselected nuclear microsatellite markers were used to assess genetic diversity, population structure, and gene flows with the aim of assembling a core collection. The dendrogram obtained utilizing the unweighted pair group method highlights the presence of homonymy and synonymy in olive tree datasets analyzed in this study. 439 different unique genotype profiles were obtained with this combination of 11 loci nSSR, representing 89.8% of the varieties analyzed. The remaining 10.2% comprises different variety pairs in which both accessions are genetically indistinguishable. Clustering analysis performed using BAPS software detected seven groups in Italian olive germplasm and gene flows were determined among identified clusters. We proposed an Italian core collection of 23 olive varieties capturing all detected alleles at microsatellites. The information collected in this study regarding the CRA-OLI ex situ collection can be used for breeding programs, for germplasm conservation, and for optimizing a strategy for the management of olive gene pools.

1. Introduction The olive (Olea europaea L. subsp. europaea var. europaea) is an important fruit species cultivated for oil and canned fruit in Italy and the Mediterranean basin. The existing ex situ collections of olive tree germplasm may valuably provide either raw material for plant breeding or plants which are directly valid for a sustainable production. With respect to the latter, we refer to those local varieties that evolved for a very long period in a location, that developed adaptative traits which are well integrated with the environmental, agronomic, cultural, and traditional features of the site, and that have been relatively recently replaced with new varieties [1]. The needs of modern agriculture, such as sustainability,

call for the cultivation of a wider range of diverse material that could better respond to the different aspects involved. Specifically, if it is necessary to obtain new varieties with a broader genetic base, capable of producing under diverse conditions and of responding to different stresses, that is, drought, pests, low fertility of the soil, and so forth, the reintroduction of old local varieties and the safeguard of traditional farming systems and landscapes can be very profitable from a socioeconomic point of views [2]. In general, the lack of information about plant genetic resources has the effect of limiting their use, restricting both the value and the usefulness of a collection even within the owning institute and among other potential users [3]. Hence, the characterisation of the germplasm conserved in a collection

2 is an essential prerequisite to a proper and wide utilization of the conserved plant material and it is the first step toward the definition of the roles that the varieties can play in sustainable production, through the direct use or in breeding programs [4, 5]. In this respect, several Mediterranean cities have promoted ex situ olive germplasm collections, including Cordoba (Spain), Marrakech (Morocco), Porquerolles (France), and Cosenza (Italy), which hosts the majority of olive varieties. Currently, on the basis of estimates from the FAO Olive Germplasm Plant Production and Protection Division, the world olive germplasm contains more than 2.600 different varieties [6], but this number could possibly have been underestimated as there is a significant lack of information regarding minor local varieties and ecotypes that are widespread in different olive-growing areas. An accurate and unambiguous identification of cultivars can be of particular importance, since different olive oils, due to their unique organoleptic and sensorial characteristics, have obtained marks of protected designation of origin (PDO) at a European level according to EC Regulation 2081/92 [7]. The main production of olive oil in Southern Italy is comprised by PDO olive oils, even though many olive cultivars with table purposes are likewise widely grown, since drupe consumption belongs to the Mediterranean diet. Over 750 million olive trees are cultivated worldwide; about 95% of them are to be found in the Mediterranean region. About 80% of the global olive oil production in 2011-2012 came from the European Union, of which 77% is concentrated in Spain, Italy, and Greece (http://ec.europa.eu/agriculture). The European Union, with about 32%, is also the major producer of world’s table olives. Even, in this case, the largest producing European countries are Spain, Greece, and Italy. Italy has about 600 olive cultivars and holds the world record for the number of cultivated varieties, representing 25% of the known world olive germplasm [8]. The Italian germplasm is large and variegated on a regional scale, because each region has gradually selected cultivars adapted to local conditions. The largest ex situ olive germplasm collection consisting of approximately 500 Italian olive varieties, and corresponding to 85% of the total Italian olive germplasm and to more than 18% of the total world olive germplasm, is maintained at the Consiglio per la Ricerca e sperimentazione per l’Agricoltura, Centro di Ricerca per l’Olivicoltura e l’Industria Olearia (CRA-OLI, Agricultural Research Council-Olive Growing and Oil Industry Research Centre) in Italy [6]. The systematic collection of Italian olive varieties for deposit into specific catalogue fields started in Italy during the 1980s. A similar international collection was initiated in 1997 by CRA-OLI of Rende, Italy. Collection included the following steps: a survey of the territory, identification, a basic characterization, and the introduction into the gene bank field. Material identified by other international scientific institutions (International Treaty on Plant Genetic Resources for Food and Agriculture-Plant Genetic Resources RGV-FAO Projects) was also included. To date, about 500 varieties have been introduced into the CRA-OLI collection (http://www.certolio.org/data-base-molecolare/). However, this wealth in terms of available biodiversity has often generated many complications in olive germplasm classification due to the lack of reference standards and

The Scientific World Journal confusion regarding the varietal names, with numerous cases of homonymy (one denomination for several genotypes) and synonymy (one genotype with several denominations) [9, 10]. It therefore appears clear how the characterization of the genetic structure is important in both the management of the olive gene pool and in understanding the role played by the domestication and subsequent crop expansion of olive trees. The identification of cultivars and accessions using molecular markers is a crucial aim of modern horticulture, with applications in breeding programs and in germplasm collection management. Traditionally olives, like other tree species, were characterized by morphological traits [11]. However, certain limitations associated with these traits have made them less popular in germplasm characterization and diversity analysis. The availability of molecular tools has provided more robust and reliable tools for germplasm characterization. In recent years, many studies about molecular characterization of germplasm in olive trees have been performed, but generally on a small number of Italian olive cultivars and without taking into account the presence of a core collection [3, 12–15]. A core collection is a subsample of a large germplasm collection that contains the minimum number of individuals that represent the whole genetic diversity and phenotypic variability of the original collection [16], essential to optimise the management and use of the large ex situ olive collections. The purpose of this study was to investigate the genetic structure of the entire Italian olive germplasm CRA-OLI collection, including all 489 accessions, using eleven nuclear microsatellite (SSRs) markers. The genetic structure of Italian olive germplasm was investigated using a model-based Bayesian clustering method to assign individuals into defined gene pools. This work is the first that takes into account such a large number of Italian olive cultivars (489 cvs) analyzed using the same set of molecular markers. This study also provides basic information for the development of core collections to maximise the representativeness of olive genetic diversity. Our results represent an essential step towards optimised conservation of olive genetic resources and subsequently for genetic association studies to detect quantitative trait loci (QTL) of adaptive and agronomic interest [17].

2. Materials and Methods 2.1. Olive Germplasm Collection. Young leaves were harvested from 489 olive trees growing in the germplasm collection of CRA-OLI, located along the Ionian coasts near MirtoCrosia (Calabrian region, Southern Italy). The analyzed olive plants are all autochthonous and representative of the seventeen regions of Italy: Abruzzo (23 varieties), Basilicata (29), Calabria (36), Campania (43), Emilia-Romagna (12), Friuli-Venezia-Giulia (3), Lazio (25), Liguria (16), Lombardy (2), Marche (19), Molise (24), Apulia (41), Sardinia (20), Sicily (70), Tuscan (101), Umbria (22), and Veneto (3) (see Table S1 in Supplementary Material available online at http://dx.doi.org/10.1155/2014/296590).

The Scientific World Journal 2.2. DNA Extraction and Microsatellites Analysis. Total genomic DNA was isolated from 100 mg fresh leaves, previously ground in liquid nitrogen, using the PureLink Genomic Plant DNA Purification Kit (Invitrogen, California, USA). DNA quality was checked on 0.9% agarose gel and the DNA concentration was estimated using NanoDrop ND2000 spectrophotometer (Thermo Scientific, Massachusetts, USA). The olive trees were genotyped at 11 nuclear SSRs, selected among those available in literature, and proven to be suitable for the characterization and identification of olive varieties in previous papers [10, 14, 18]: four SSRs (GAPU59, GAPU71A, GAPU71B, and GAPU103A) by Carriero et al. [19], five (UDO01, UDO03 UDO12, UDO28, and UDO39) by Cipriani et al. [20], and two (DCA9 and DCA18) by Sefc et al. [21]. The PCR was conducted in a final volume of 25 𝜇L containing 25 ng of DNA, 10 mM Tris-HCl pH 8.0, 1.5 mM MgCl2 , 0.2 mM dNTPs, 0.25 𝜇M forward and reverse primers, and 0.05 units of Taq DNA polymerase (Invitrogen, California, USA) as reported in Muzzalupo et al. [22]. SSR amplification was carried out as described by Muzzalupo et al. [14]. The amplification products were analyzed by means of a 2100 Bio-Analyzer instrument (Agilent Technologies, Waldbronn, Germany) with the 2100 BioSizing software (version A.02.12) using DNA 500 LabChip kit. To assign the correct size to alleles, most alleles of the selected loci were sequenced. The allele sequencing was carried out as described by Baldoni et al. [18].

3 of the Italian olive germplasm, a model-based analysis was performed using BAPS 5.3 [27]. This program uses both a nonspatial and spatial Bayesian clustering algorithm assignment to determine the number of genetically distinct populations present in a sample based on allele frequencies. We conducted admixture and mixture analysis on olive varieties distributed at the regional level and using the nonspatial model. BAPS was run setting 1000 as the number of interactions used to estimate the admixture coefficients for the genotypes, 200 as the number of reference individuals from each genotype, and 10 as the number of interactions used to estimate the admixture coefficients for the reference individuals, reanalyzing and comparing our data set also using smaller (5) and higher (20) values. In addition, an Analysis of Molecular Variance (AMOVA) [28, 29] was performed to estimate levels of genetic differentiation by computing ΦPT , 𝐹ST , and 𝑅ST estimators among BAPS groups identified in this study. Statistical significance of all the ΦPT , 𝐹ST , and 𝑅ST estimators were tested using 10,000 permutations. Finally, we used the GraphViz 2.28 package installed in BAPS 5.3, to estimate and draw the gene flows among the clusters identified. In the graph, gene flows were shown by weighted arrows, so that the weights relating to the amounts of ancestry in the source cluster were assigned to the target cluster. This step was performed using the result file from the dataset admixture analysis and by setting the threshold for the significance of 𝑃 values of the admixture estimates to 0.05.

2.3. Data Analysis 2.3.1. Genetic Diversity and Multivariate Analysis. Number of alleles (𝑁𝑎), effective number of alleles (𝑁𝑒), observed (𝐻𝑜) and expected (𝐻𝑒) heterozygosity, and fixation index (𝐹) were computed with GenALEx version 6.5 software [23]. The alleles detected for each microsatellite were recorded into a data matrix of presence (1) and absence (0) of bands (each allele representing a band). Genetical distance based on the Nei coefficient and genetic similarity based on the Simple Matching (SM) coefficient among 489 olive varieties were estimated using the NTSYSpc program version 2.02 [24]. Finally, a tree was inferred using the unweighted pair group method using an Arithmetic average (UPGMA) clustering algorithm to highlight the presence or absence of synonymies in the olive varieties data set analyzed in this study. In addition, the dendrogram was tested by bootstrapping to determine the confidence limits and using WinBoot program [25]. The frequency of null alleles was estimated per locus and per region, using the software FreeNA [26]. Principal Coordinate Analysis (PCoA), also available in version 6.5 of the GenALEx program, was conducted using Nei’s unbiased genetic distance pairwise population matrix to determine whether observed patterns in molecular data support the partitioning of the olive tree samples into specific groupings. 2.3.2. Bayesian Model-Based Clustering Analysis, Molecular Variance, and Gene Flows. To study the genetic structure

2.3.3. Core Collection Sampling. The Maximisation strategy [30–32] implemented in the COREFINDER software [33] was used to generate core olive collection that maximised the number of observed alleles in our nuclear dataset. The M-strategy consists in detecting the best sample size that captures 100% of the genetic diversity present within the entire germplasm collection. The algorithm is based on the Set-Covering (NP-complete) problem. The procedure is a Las Vegas style randomized algorithm: an iteration number is provided by the user, and the algorithm, starting from a random initial set, uses a greedy strategy to search for an accession “A” providing a better overall genetic diversity than some accession “B” belonging to the current core collection. In such a hypothesis, “A” is included and “B” is excluded from the collection. The greedy step is performed exhaustively and each iteration starts with a different initial random set, thereby reducing the probability of ending in a local maximum. In our COREFINDER analysis, the algorithm parameters were set on 100 and 1.000.000 for interations and random seed, respectively.

3. Results 3.1. Genetic Diversity. Eleven published primer pairs flanking nuclear microsatellites were employed to investigate the level of genetic variation among the 489 Italian olive varieties analyzed in this study and present in the olive germplasm collection of the CRA-OLI. A total of 84 alleles over 11 loci were detected, ranging from 3 at UDO01 locus to 12 alleles at

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Table 1: Genetic diversity parameters estimated for the SSR loci in the 489 olive varieties. For each locus, the number of alleles detected (𝑁𝑎), the effective number of alleles (𝑁𝑒), the observed (𝐻𝑜) and expected (𝐻𝑒) heterozygosity, the fixation index (𝐹), and the frequency of null allele (𝑁𝑢) are reported. Locus GAPU59 GAPU71A GAPU71B GAPU103A UDO01 UDO03 UDO12 UDO28 UDO39 DCA09 DCA18 Mean SE

𝑁𝑎 5 9 5 8 3 6 6 9 12 12 9 7.6 —

𝑁𝑒 4.0 3.1 3.6 5.7 2.3 3.2 3.9 5.6 5.3 6.7 4.0 4.3 0.090

𝐻𝑜 0.632 0.578 0.885 0.786 0.074 0.083 0.828 0.793 0.330 0.903 0.760 0.605 0.024

𝐻𝑒 0.637 0.602 0.684 0.755 0.504 0.543 0.671 0.735 0.704 0.785 0.686 0.664 0.010

𝐹 0.013 0.018 −0.300 −0.068 0.860 0.857 −0.260 −0.106 0.544 −0.153 −0.116 0.114 0.035

𝑁𝑢 0.042 0.051 0.002 0.034 0.297 0.295 0.001 0.036 0.214 0.002 0.008 — —

The presence of null alleles are indicated in bold.

both UDO39 and DCA9 loci. The shortest allele among the 11 polymorphic loci was allele 108 base pairs (bp) at UDO39, whereas the longest was 259 bp at GAPU71A (Table S2). The most common size variant, namely, the allele 214 bp at locus GAPU71A, was found with a frequency of 0.485 and showed the highest value in the varieties within Molise region (0.813). Three out of 84 alleles were considered private alleles, since they were present only in “Cellina di Nard`o” (allele 228 bp at locus GAPU71A), in “Arancino,” “Ciliegino,” “Emilia,” “Grappolo,” “Gremignolo,” “Gremignolo di Bolgheri,” “Lastrino,” “Lazzera reale,” “Salicino,” “Santa Caterina,” “Borgiona,” and “Corniolo” (allele 156 bp at locus UDO12), and in “Filare” (allele 184 bp at locus DCA9) with a frequency 0.20 of null allele frequency have been considered as a threshold over which a significant underestimation of 𝐻𝑒 due to null alleles can be found. Frequencies lower than 0.20 were obtained for all loci for each of the sampled varieties, except for UDO01 (0.297), UDO03 (0.295), and UDO39 (0.214) loci, where null allele frequencies higher than 0.20 were found (Table 1). For this reason, these loci were eliminated from further analysis. 3.2. Characterisation of Olive Accessions. The dendrogram (Figure S1) obtained utilizing the UPGMA method that elaborates a matrix of similarity obtained using NTSYSpc program version 2.02 [24] was tested using WinBoot program [25] and highlights the presence or absence of mislabeling, redundancies, homonymy, and synonymy in olive tree datasets analyzed in this study. 439 different unique genotype

profiles were obtained with this combination of 11 loci, being able to identify about 89.8% of the varieties analyzed showing unique profiles. The remaining 10.2% is comprised of different variety pairs in which both accessions are genetically indistinguishable one from another, which has the potential to represent cases of synonymy (Table 2). Synonyms included cultivars with the same profile for all SSR examined and cultivar pairs differing from each other for one or two alleles [14, 34, 35]. Ten different olive cultivar pairs or groups are genetically indistinguishable from one another (Table 2). Many of these possible cases of synonymy are in agreement with previous studies based on morphological descriptors and molecular marker systems [10, 14, 15, 22, 36]; others were encountered for the first time. Parent-offspring relations were found for “Giarraffa” and “Pizzo di Corvo,” “Nera di Oliena” and “Paschixedda,” “Cazzarella” and “Sperone di Gallo,” “Grossa di Venafro” and “Paesana Nera,” “Paesana Bianca” and “Rosciola di Rotello,” “Racemo 1” and “Coratina,” “Rossina” and “Selvatico,” and “Toccolana” and “Olivetta nera,” and these eight cultivar pairs differ by only one allele (Table 2). Parent-offspring relations were found for “Ascolana dura” and “Ascolana semitenera,” “Dolce di Andria” and “Termite di Bitetto,” “Paschixedda” and “Terza Piccola,” “Gentile nera di Colletorto” and “Noccioluta,” “Ginestrino” and “Maurino 2,” “Ginestrino” and “Maurino 4,” “Maurino 2” and “Maurino 4,” “Leccio del Corno 2” and “Piangente 3,” “Nerba” and “Olivo di Castiglione,” and “Nostrale di Fiano Romano” and “Raza.” These ten cultivar pairs differ by two alleles. Furthermore, 22 cases of homonymy were identified. These 22 cases of homonymy can be divided into two groups according to the number of different alleles. The first group is represented by plants that have a number of different alleles less than ten. The list of homologies is shown in Table 2. A special case is that presented in the group of “Leccino,” “Moraiolo,” “Pendolino,” “Maurino,” “Nostrana di Brisighella,” and “San Felice Acquasparta” which, in previous

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Table 2: Potential cases of mislabelling, redundancies, homonymy, and synonymy identified by microsatellite fingerprinting on 489 Italian olive varieties. Possible cases of synonyms “Carolea Cefaly”—“Carolea Mirto” “Carolea Cetraro”—“Carolea Rossi” “Cima di Mola”—“Ogliarola salentina” “Leccino Dwarf ”—“Leccino Minerva” (plant 1)—“Leccino Pisa” (plants 4 and 9) “Majorca”—“Manna” “Mele”—“Nolca” “Nera di Gonnos”—“Tonda di Cagliari” “Nera di Oliena”—“Terza piccola” “Nera di Villacidro”—“Terza grande” “Ogliarola del Bradano” (plant 1)—“Taggiasca”—“Casaliva”—“Ogliarola barese”—“Ogliarola garganica”—“Correggiolo”—“Correggiolo Montegridolfo”—“Correggiolo Pallese”—“Frantoio” (plants 1/7 − 10)—“Frantoio FC”—“Frantoio Villa Verrucchio”—“Frantoio Montegridolfo”

Genotype 1 2 3 4 5 6 7 8 9 10

Plants with the same genotypes (G) “Arnasca” (plants 1, 2, 3, and 4) “Buscionetto” (plants 2 and 3) “Canino” (plants 1 and 2) “Coratina” (plants 1, 2 and 3) “Frantoio” (plants 8 and 9) “Gentile di Larino” (plants 1 and 5) “Iacona” (plants 1 and 2) “Leccino” (plants 3 and 4) “Leccino” (plants 6, 7, 8, and 9) “Maiatica di Ferrantina” (plants 1 and 2) “Moresca” (plants 1 and 2) “Nocellara del Belice” (plants 1 and 2) “Nocellara Nissena” (plants 1 and 2) “Pirunara” (plants 1 and 2) “Rotondella di sanza” (plants 1 and 2) “San Felice Acquasparta” (plants 4 and 5) Possible cases of synonyms: plants with different genotypes (one allele) “Cazzarella” and “Sperone di gallo” “Giarraffa” and “Pizzo di corvo” “Grossa di Venafro” and “Paesana nera” “Leccino 1” and “Leccino 10” “Maurino” (plants 1 and 3) “Moraiolo” (plants 1 and 3) “Moraiolo” (plants 2 and 4) “Nera di Oliena” and “Paschixedda” “Nostrana di Brisighella” (plants 2 and 4) “Paesana bianca” and “Rosciola di Rotello” “Pendolino” (plants 2 and 5) “Racemo 1” and “Coratina” (plants 1, 2, and 3) “Rossina” and “Selvatico” “Toccolana” and “Olivetta nera”

“Arnasca G” “Buscionetto G” “Canino G” “Coratina G” “Frantoio G” “Gentile di Larino G” “Iacona G” “Leccino G1” “Leccino G2” “Maiatica Ferrantina G” “Moresca G” “Nocellara del Belice G” “Nocellara Nissena G” “Pirunara G” “Rotondella di sanza G” “San Felice Acquasparta G”

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Table 2: Continued. Possible cases of synonyms: plants with different genotypes (two alleles) “Ascolana dura” and “Ascolana semitenera” “Dolce di Andria” and “Termite di Bitetto” “Gentile nera di Colletorto” and “Noccioluta” “Ginestrino” and “Maurino” (plants 2 or 4) “Leccio del Corno 2” and “Piangente 3” “Maurino” (plants 2 and 4) “Moraiolo” (plants 4 and 5) “Nerba” and “Olivo di Castiglione” “Nostrale di Fiano Romano” and “Raza” “Paschixedda” and “Terza piccola” “Pendolino” (plants 1 and 5) “Pendolino” (plants 3 and 4) Homonyms: plants with different genotypes (from three to nine alleles) “Buscionetto 1” and “Buscionetto G” “Cucca” (plants 1 and 2) “Erbano” (plants 1 and 2) “Faresana” (plants 1 and 2) “Leccino G1” and “Leccino G2” “Leccino 2” and “Leccino G1” “Leccino” (plants 2 and 10) “Leccino Minerva” (plants 1 and 2) “Moraiolo” (plants 1 and 5) “Nostrana di Brisighella” (plants 1 and 3) “Ogliarola messinese” (plants 1 and 2) “Pendolino” (plants 1 and 3) “Piangente” (plants 1 and 3) “Pizzo di corvo” (plants 1 and 2) “Razzo” (plants 1 and 2) “Turdunazza antimosca” (plants 1 and 2) “Vallanella” (plants 1 and 2) Homonyms: plants with different genotypes (ten or more alleles) “Arnasca 4” and “Arnasca G” “Giarraffa” (plants 1 and 2) “Leccio del Corno” (plants 1 and 2) “Minuta” (plants 1 and 2) “Minuta” (plants1 and 3) “Minuta” (plants 2 and 3) “Ogliarola del Bradano” (plants 1 and 2) “Piangente” (plants 1 and 2) “Racioppa” (plants 1 and 2) “Romanella” (plants 1 and 2) “San Felice Acquasparta” (plants 1 and 2) “San Felice Acquasparta” (plants 2 and 3) “San Felice Acquasparta 3” and “San Felice Acquasparta G” “Sargano” (plants 1 and 2)

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3.3. Genetic Structure of Italian Olive Genotypes. Genetic structure was tested using two different approaches. First, the PCoA, performed on Nei’s unbiased genetic distance matrix and, based on 62 different size variants, showed that the 439 olive varieties were separated into five main groups (Figure 1). Group I contains olive varieties of Molise (23), Tuscany (79), Abruzzo (23), Basilicata (28), Apulia (35), and Sicily (64). Group II comprises Calabria (34) and Veneto (3) varieties. Group III include the olive varieties present in the regions of Lombardy (1), Lazio (24), Liguria (12), Marche (19), Umbria (21), and Emilia-Romagna (12), while Group IV contains varieties of Campania (42) and Sardinia (16). Group V only contains varieties from Friuli-Venezia-Giulia (3) region and is slightly more distant from genetic groups previously described. In the PCoA analysis, the first two principal axes explain a total of 68.9% of unbiased genetic distance, with 54.4% and 14.5% for coordinates 1 and 2, respectively (Figure 1). Bayesian clustering algorithms, such as those implemented in BAPS 5.3 program, were used for inferring olive Italian germplasm structure [27]. Seven genetic clusters were identified with a more specific distribution of genotypes than PCoA analysis. The identified clusters are represented in Figure 2. Our analysis showed that all the olive varieties present in Northern and Central Italy were grouped in a single genetic cluster, with the exception of the varieties present in the regions of Tuscany, Abruzzo, and Molise that were grouped into two separate clusters (Figure 2). The BAPS analysis also revealed how the varieties distributed in the remaining regions of Southern Italy (Campania, Apulia, Calabria, and Basilicata), as well as those present in the two major islands of Sicily and Sardinia, are grouped into four distinct genetic clusters (Figure 2). Furthermore, BAPS analysis showed high levels of 𝐻𝑜 in each genetic cluster identified and ranged from 0.680 to 0.852

SAR CAM Coordinate 2

work, had been regarded as polyclonal varieties [22, 37]. In fact, the “Leccino” group is represented by 10 plants that clustered forming subgroups with microsatellite profiles that differ by a minimum of one allele to a maximum of eight alleles. A similar pattern was observed in the case of the group of “Moraiolo” (4 plants), of “Maurino” (4), of “Pendolino” (5), and “Nostrale di Brisighella” (4) that show differences in the profile microsatellite, from 1 to 3 alleles, from 1 to 2 alleles, and from 3 to 9 alleles, respectively. Finally, the “San Felice Acquasparta” denomination is represented by five plants that do not cluster. Only plants 4 and 5 formed subgroups with the same SSR profiles; however, they were differentiated from plants 1, 2, and 3 by a minimum of nine to a maximum of ten alleles. Finally, SSR analysis allowed the classification of the CRA-OLI olive germplasm into 439 unique molecular profiles corresponding to well-defined genotypes, whereas, for the remaining molecular profiles, they reveal the presence of accessions considered as clones or possible synonyms of the same genotype.

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TUS MOL ABR

LOM

BAS APL SIC CAL

LIG LAZ MAR

UMB EMR

VEN

FVG

Coordinate 1

Figure 1: Principal Coordinates Analysis (PCoA) plot of the olive varieties based on the first two principal coordinates (coord. 1 = 54.43% and coord. 2 = 14.52%). Legend: ABR: Abruzzo, APL: Apulia, BAS: Basilicata, CAL: Calabria, CAM: Campania, EMR: EmiliaRomagna, FVG: Friuli-Venezia-Giulia, LAZ: Lazio, LIG: Liguria, LOM: Lombardy, MAR: Marche, MOL: Molise, SAR: Sardinia, SIC: Sicily, TUS: Tuscany, UMB: Umbria, and VEN: Veneto.

with a mean of 0.755, than 𝐻𝑒 levels (mean 0.724) producing a negative value of 𝐹 (mean −0.045) (Table 3). In addition, the AMOVA analysis revealed comparable values of 𝐹ST and 𝑅ST estimators among BAPS groups (𝐹ST = 5.7 and 𝑅ST = 5.2), with a higher ΦPT estimator value (ΦPT = 10.9) than those reported in literature for Olea [37]. All of the AMOVA analysis conducted as part of this study showed that most of the diversity being expressed within BAPS groups identified for all the estimators is considered (Table 4). 3.4. Gene Flows. The analysis of the CRA-OLI Italian olive germplasm collection performed with a Bayesian clustering software, demonstrated a good network of gene flows between the clusters identified in this study. This analysis also revealed how only the clusters that grouped the olive varieties present in Sicily (64), Tuscany (79), Sardinia (16), Apulia (35), Basilicata (28), and Calabria (34) and those included in Northern and Central Italy were more susceptible to the identified gene flows, with a consequent transfer of genetic material (Figure 3). The analysis of gene flows showed that clusters 3 (Sardinia), 6 (Emilia-Romagna, FriuliVenezia-Giulia, Lazio, Liguria, Lombardy, Marche, Umbria, and Veneto), and 7 (Basilicata, Calabria, and Apulia) were characterized by a higher level of output gene flow, while only clusters 6 and 7 had a very high level of input genetic material transfer. On the other hand, this analysis showed that, only in cluster 1 (Abruzzo and Molise), no input gene flows with other clusters seemed to occur. 3.5. Core Collection. A core collection was herein assembled for Italian olive germplasm, aiming to represent the entire genetic diversity identified in this study. The COREFINDER analysis based on M-strategy showed that, for Italian olive germplasm, 100% of the SSR alleles found in this study could be represented by a core collection of 23 accessions (Figure 4 and Table S3). In addition, our COREFINDER analysis highlighted that 39% of the entire core collection

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56 7 89 10 11

12

13

14

15

16 17

1 (Abruzzo), 11 (Molise) 2 (Basilicata), 3 (Calabria), 12 (Apulia) 4 (Campania) 5 (Emilia-Romagna), 6 (Friuli Venezia Giulia), 7 (Lazio), 8 (Liguria), 9 (Lombardy), 10 (Marche); 16 (Umbria), 17 (Veneto) 13 (Sardinia) 14 (Sicily) 15 (Tuscan)

Figure 2: BAPS analysis on the 439 Italian olive varieties. In the graph, each colour represents a population group based on allele frequency. Vertical bars represent each olive variety analyzed in this study and bars are divided into several colours when there is evidence of admixture. Table 3: Genetic diversity parameters at SSR loci estimated in BAPS groups identified in this study. For each cluster, the observed heterozygosity (𝐻𝑜), the expected heterozygosity (𝐻𝑒), and the fixation index (𝐹) are reported. BAPS clusters Cluster 1 Cluster 2 Cluster 3 Cluster 4 Cluster 5 Cluster 6 Cluster 7 Mean SE

𝐻𝑜 0.734 0.725 0.852 0.762 0.680 0.796 0.736 0.755 0.020

was represented by the olive varieties grouped in cluster 1 (Abruzzo and Molise) identified by BAPS analysis. Other BAPS clusters contribute to the core collection at smaller percentages: cluster 6 (26%), cluster 2 (13%), clusters 4 and 7 (9%), and cluster 3 (3%). Overall, the core collection identified in this study represents 5.2% of the CRA-OLI olive germplasm collection.

4. Discussion The results by SSR analysis of CRA-OLI Italian olive germplasm collection show abundant allelic variation over 11 loci and high overall genetic diversity, confirming that SSR markers can be effectively used to genotype a germplasm collection. Four of these loci were included in the best consensus set of SSR markers [18] that has already been used for genetic structure studies [14]. It is currently well known how mating systems play a key role in determining the structure of genetic diversity in natural and domesticated genotypes. This is especially true for olive trees that have been clonally propagated since ancient times. This claim was also confirmed in our study by clustering analysis, NTSYS, and PCoA, performed on our

𝐻𝑒 0.700 0.737 0.710 0.750 0.665 0.763 0.744 0.724 0.012

𝐹 −0.062 0.018 −0.208 −0.017 −0.017 −0.046 0.015 −0.045 0.069

dataset. For NTSYS analysis, the results demonstrated the presence of synonyms and homonyms among the different varieties in the Italian olive germplasm which are partially comparable to those reported in literature. Homonymy and synonymy characterization is essential in order to avoid genotype redundancy and to maximize genetic diversity in the Italian olive germplasm collection. Additionally, the PCoA analysis showed a clear grouping of the olive Italian varieties into five main clusters, broadly confirming previously reported results [12–14]. Bayesian analysis (BAPS) further provided support for the existence of genetic structure in CRA-OLI germplasm collection and separated the Italian olive varieties into seven main clusters. The Bayesian model-based analysis highlights the real structure and distribution of Italian olive germplasm gene pools, separating the major genetic cluster 6 that grouped olive varieties in Northern and Central Italy, from the other six gene pools found. In addition, BAPS analysis results show the genetic relationship, represented by gene flows, among the seven clusters identified, confirming that the current gene pools and distribution of Italian olive germplasm are due to geographic and cultural aspects mainly involving human activity in the past.

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Table 4: AMOVA analysis for the partitioning of SSR variation of olive varieties among and within BAPS groups identified in this study. Estimators ΦPT ΦPT 𝐹ST 𝑅ST

Source of variation Among groups Within groups Among groups Within groups Among groups Within groups

df 6 432 6 871 6 871

Variance components 0.709 5.814 0.177 2.944 68.279 1234.244

𝑃 value

Percentage total variance 10.9 89.1 5.7 94.3 5.2 94.8

𝑃 < 0.001 𝑃 < 0.001 𝑃 < 0.001

1

Cluster 1

0.03

0.95

Cluster 5

0.0087

0.024

0.0017 0.00031

0.015

Cluster 2

0.013

0.013

0.00031

0.002

0.94

0.0037

0.031

0.00063

Cluster 3

0.021

0.94

0.0013

0.00031

0.028

0.0047 0.042 0.052

Cluster 7

0.016 0.0076

0.96 0.025

0.012

0.00018 Cluster 6

0.00072

0.84

0.015 0.011 0.0062 Cluster 4

0.98

Figure 3: Main gene flows between clusters identified in Italian olive germplasm collection of CRA-OLI. In the graph, gene flows were shown by weighted arrows, so that the weights relative amounts of ancestry in the source cluster were assigned to the target cluster.

Moreover, the AMOVA results surprisingly revealed a higher ΦPT estimator value (ΦPT = 10.9) than those reported in literature for Olea europaea [37], showing a good level of genetic differentiation distributed at a genetic cluster level among Italian olive varieties in the CRA-OLI collection. This result was confirmed by higher levels of 𝐻𝑜 and 𝐻𝑒 detected in BAPS clusters, with a mean negative value of 𝐹 that clearly highlights the good levels of genetic diversity,

maintained costant in each BAPS group identified in this study. The final aim of the study was to construct a valid core collection for cultivated olives in the CRA-OLI germplasm collection, sampling the minimum number of entries that maximize the representativeness of allelic diversity. The core collection that we proposed in this study consists of 23 varieties that capture 100% of the base collection. It was found

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10

100 80 60 40 20 0

𝐹-statistics Genetic Analysis in Excel Expected heterozygosity Observed heterozygosity Observed number of alleles Effective number of alleles Numerical Taxonomy and Multivariate Analysis System 𝑁𝑢: Null allele nSSR: Nuclear Simple Sequence Repeat PCoA: Principal Coordinates Analysis 𝑅-statistics 𝑅ST : SM: Simple matching SSRs: Simple sequence repeats UPGMA: Unweighted pair group method using an Arithmetic average PhiPT. ΦPT :

𝐹ST : GenAlEx: 𝐻𝑒: 𝐻𝑜: 𝑁𝑎: 𝑁𝑒: NTSYSpc:

120

1

3

5

7

9

11

13

15

17

19

21

23 25

Number of accessions of the core

Figure 4: Genetic diversity as a function of the number of accessions included in the Italian olive germplasm core collection.

that only a small number of olive varieties, compared with other values reported in literature for Olea [3], are necessary to represent the molecular diversity revealed in this study. These results are probably due to the percentage of cluster 1 representativeness (39%) within the core collection, further confirming the genetic peculiarity of this cluster, already highlighted by gene flow analysis. Nevertheless, high levels of heterozygosity observed in Italian olive germplasm may contribute to reducing the size of the core collection [33]. Van Hintum [38] suggested that the sampling proportion should vary between 5 and 20% of the base collection, representing at least 70% of overall genetic diversity. The core collections for Italian olive germplasm proposed here represent 100% of the molecular diversity found in this study, with the number of varieties accounting for 5.2% of the CRA-OLI germplasm collection. Finally, the result of our BAPS analysis supports both NTSYS and PCoA results, demonstrating how all olive accessions analyzed in this study and maintained in CRAOLI ex situ collection could be considered representative of Italian olive germplasm because each genetic group defined in BAPS reflects geographic distribution and confirms that the Italian olive germplasm is a peculiar gene pool present in the Mediterranean basin. Concluding, the use of molecular markers, like microsatellite, is imperative to build a database for cultivar analysis, for the traceability of processed food, and for the appropriate management of olive germplasm collections. Moreover, the results presented here regarding clustering and core collection are extremely useful for the selections of parents to be used for breeding programs and thus ensuring an optimal management of the CRA-OLI Italian olive germplasm collection. This work is the first study with such a large number of Italian olive varieties analyzed using the same set of molecular markers which allowed characterisation of the genetic structure and identification of a core collection in the largest Italian ex situ germplasm collection.

Abbreviations AMOVA: Analysis of Molecular Variance BAPS: Bayesian Analysis of Population Structure 𝐹: Fixation index

Conflict of Interests The authors declare that there is no conflict of interests regarding the publication of this paper.

Acknowledgments The authors are grateful to S. Bonavita for his help with the statistical analysis. This work was supported by the Ministry of Economic Development, (Certificazione della composizione varietale, dell’origine geografica e dell’assenza di prodotti di sintesi negli oli extravergini di oliva - CERTOLIO project) and by the Ministry of Agriculture, Food and Forestry (Trattato internazionale FAO sulle risorse genetiche vegetali, III triennalit`a 2010–2013, RGV-FAO project).

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Table S1 List of the Olea europaea L.varieties present in the CRA-OLI germplasm collection and analyzed by microsatellite markers. The Italian olive varieties included in the core collection are indicated in bold. N. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46

Varieties Abunara Agristigna Aitana Albatro Allora Amati Americano Annarea Arancino Arnasca 1 Arnasca 2 Arnasca 3 Arnasca 4 Arnasca 5 Arnasca 6 Ascolana dura Ascolana semitenera Ascolana tenera Augellina Aurina Barilari Bianchella Bianchera Biancolella Biancolilla Borgese Borgiona Bosana Bottone di Gallo 1 Bottone di Gallo 2 Bottoni di Gallo Brandofino Buga Buscionetto 1 Buscionetto 2 Buscionetto 3 Cacaredda Cacaridduni Cacazzara Caiazzana Calatina Cammarotana Canino 1 Canino 2 Capolga Caprina di Casalanguida

Regions Sicily Calabria Sicily Toscany Toscany Lazio Toscany Basilicata Toscany Liguria Liguria Liguria Liguria Liguria Liguria Marche Marche Marche Basilicata Molise Sicily Umbria Friuli Venezia Giulia Campania Sicily Calabria Umbria Sardinia Sicily Sicily Molise Sicily Friuli Venezia Giulia Sicily Sicily Sicily Sicily Sicily Sicily Campania Sicily Campania Lazio Lazio Marche Abruzzo

Code B # 205 C1 # 319 B # 206 C1 # 270 C1 # 271 C1 # 338 A # 139 B # 262 C1 # 272 A # 46A A # 46B A # 46C A # 46D A # 46E A # 46 B # 229 B # 230 A # 53 A # 10 B # 184 C1 # 346 A # 128 A # 159 C2 # 358 C1 # 347 A # 18 B # 180 A # 71 C1 # 308A C1 # 308B A # 174 A # 78 C1 # 337 A # 79A A # 79B A # A32 B # 207 B # 208 C2 # 383 B # 240 A # 124 C1 # 324 A # 32A A # 32B B # 231 B # 260

47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97

Caprina Vastese Carbona Carboncella Carboncella Campana Carboncella Pianacce A Carboncella Pianacce B Carbonchio Carbuncion Carbuncion di Carpinenga Cariasina Cariola Carmelitana Carolea Cefaly Carolea Cetraro Carolea Mirto Carolea Rende Carolea Rossi Carpellese Carpinella Carpinetana Casaliva Castiglionese Castricianella Rapparina Cavalieri Cazzarella Cazzinicchio Cellacchia Cellina di Nardo' Cellina di Rotello Cerasa di Montenero Cerasuola Ciciarello Cicinella Ciliegino Cima di Melfi Cima di Mola Colombina Coratina 1 Coratina 2 Coratina 3 Cornacciola Corneglia Cornia Corniola Corniolo Cornula Coroncina Correggiolo Correggiolo di Montegridolfo Correggiolo di Pallese Corsicana da mensa

Abruzzo Friuli Venezia Giulia Lazio Campania Lazio Lazio Abruzzo Emilia Romagna Emilia Romagna Sardinia Calabria Apulia Calabria Calabria Calabria Calabria Calabria Campania Basilicata Abruzzo Lombardy Abruzzo Sicily Sicily Molise Apulia Lazio Apulia Molise Molise Sicily Calabria Campania Toscany Basilicata Apulia Emilia Romagna Apulia Apulia Apulia Basilicata Campania Campania Calabria Umbria Apulia Marche Toscany Toscany Toscany Sardinia

B # 261 C2 # 367 A # 33 C2 # 359 B # A5 B # A6 B # 216 C1 # 334 C2 # 397 A # 72 C2 # 400 A # 120 B # A7 B # A38 A # 19 R P1F14 B # A8 C1 # 325 C2 # 390 B # 204 A # 51 A#1 B # 209 B # 182 C2 # 369 A # 121 C1 # 339 A # 55 B # 185 C2 # 370 A # 80 A # 115 C1 # 326 C1 # 273 A # 11 A # 56 B # 198 C2 # A2-057 A # A24 A # 57 C2 # 391 B # 241 B # 242 B # 192 A # 170 R P1F08 A # 103 A # 126 A # A30 C1 # 274 C1 # 311

98 99 100 101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120 121 122 123 124 125 126 127 128 129 130 131 132 133 134 135 136 137 138 139 140 141 142 143 144 145 146 147 148

Corsicana da olio Craputea Crastu Crognolo CSM CSS-02-Minerva Cucca 1 Cucca 2 Curatora Demoniata Dolce Agogia Dolce di Andria Dolce di Cassano Dolce di Cerchiara Dolce di Rossano Don Carlo Donna Francesca Dritta Dritta Loreto Emilia Erbano 1 Erbano 2 Faresana Faresana Fasolina Fasolona Favarol Fecciaro Feglina Femminella di Torraca Filare Fosco Frangivento Frantoio 1 Frantoio 10 Frantoio 2 Frantoio 3 Frantoio 4 Frantoio 5 Frantoio 6 Frantoio 7 Frantoio 8 Frantoio 9 Frantoio di Montegridolfo Frantoio di Villa Verrucchio Frantoio FC Fs17 Gaggiolo Gentile dell'Aquila Gentile di Chieti Gentile di Larino 1

Sardinia Emilia Romagna Sicily Lazio Toscany Toscany Toscany Toscany Campania Sicily Umbria Apulia Apulia Calabria Calabria Umbria Apulia Abruzzo Marche Toscany Sicily Sicily Basilicata Basilicata Basilicata Basilicata Veneto Umbria Liguria Campania Toscany Lazio Apulia Toscany Toscany Toscany Toscany Toscany Toscany Toscany Toscany Toscany Toscany Toscany Toscany Toscany Umbria Lombardy Abruzzo Abruzzo Molise

A # 73 C1 # 264 C1 # 348 C1 # 340 B # A3 C1 # A355 A # 127 C1 # A1-127 C2 # 366 C2 # 384 A # 97 B # 196 A # 122 C1 # 320 A # 26 C2 # 389 C2 # 403 A#4 B # 232 C1 # 276 C1 # 349A C1 # 349B B # 202 C1 # A1-202 C1 # 265 C1 # 266 A # 119 A # 171 A # 47 B # 243 C1 # 277 B # 227 A # 58 C2 # A8-092 B # A11 A # 92 C2 # A3-092 C2 # A1-092 A # A6-092 B # A9 B # A10A-092 B # A10B-092 C2 # A7-092 A # A28 A # A29 B # A12 A # 154 A # 52 B # 217 A#5 A # 166A

149 150 151 152 153 154 155 156 157 158 159 160 161 162 163 164 165 166 167 168 169 170 171 172 173 174 175 176 177 178 179 180 181 182 183 184 185 186 187 188 189 190 191 192 193 194 195 196 197 198 199

Gentile di Larino 5 Gentile nera di Colletorto Ghiacciolo Ghiacciora Ghiandaro Ghiannara Giardino Giarfara Giarraffa 1 Giarraffa 2 Ginestrino Giusta Gnagnaro Grappolo Grappuda Gremignolo Gremignolo di Bolgheri Grignan Grossa di Cassano Grossa di Gerace Grossa di Spagna Grossa di Venafro Grossale I 77 Iacona 1 Iacona 2 Imperiale Intosso Itrana Larcianese Lastrino Laurina Lavagnina Lazzera Reale Lea Leccino 1 Leccino 10 Leccino 2 Leccino 3 Leccino 4 Leccino 5 Leccino 6 Leccino 7 Leccino 8 Leccino 9 Leccino Compatto Leccino Dwarf Leccino Minerva 1 Leccino Minerva 2 Leccino Pisa 4 Leccino Pisa 9

Molise Molise Calabria Calabria Abruzzo Basilicata Liguria Sicily Sicily Sicily Toscany Basilicata Molise Toscany Emilia Romagna Toscany Toscany Veneto Calabria Calabria Apulia Molise Campania Umbria Sicily Sicily Calabria Abruzzo Lazio Toscany Toscany Marche Liguria Toscany Marche Toscany Toscany Toscany Toscany Toscany Toscany Toscany Toscany Toscany Toscany Toscany Toscany Toscany Toscany Toscany Toscany

A # 166B B # 190 C2 # 398 R P1F10 A#2 B # 263 A # 121 B # 210 A # 81A A # 81B C1 # 279 B # 223 B # 187 C1 # 238 C2 # 388 C1 # 281 C1 # 282 A # 157 A # 20 A # 21 A # 59 B # 188 B # 244 B # 179 C1 # 350A C1 # 350B A # 158 A#6 A # 34 C1 # 284 C1 # 285 A # 151 A # 164 C1 # 286 A # 104 C2 #A9-093 B # A20-093 C1 # A8-093 C1 # A7-093 A # 93 A # A27 A # A6-093 A # A5-093 A # A4-093 A # A3-093 B # A17 B # A18 A # A26A A # A26A C2 # 093A C2 # 093B

200 201 202 203 204 205 206 207 208 209 210 211 212 213 214 215 216 217 218 219 220 221 222 223 224 225 226 227 228 229 230 231 232 233 234 235 236 237 238 239 240 241 242 243 244 245 246 247 248 249 250

Leccio del Corno 1 Leccio del Corno 2 Leccio Marammano 1 Leccio Maremmano 2 Leccione Lezze Lumiaru Mafra Mafra di Cerchiara Maglianese Maiatica di Ferrantina 1 Maiatica di Ferrantina 2 Majorca Mandanici Manna Mantonica Marchiaio Marina Marinese Marzio Maurino 1 Maurino 2 Maurino 3 Maurino 4 Mele Mignola Mignolo Mignolo Cerretano Minna di Vacca Minuta 1 Minuta 2 Minuta3 Minutella Monaca Mora Moraiolo 1 Moraiolo 2 Moraiolo 3 Moraiolo 4 Moraiolo 5 Moraiolo T. Corsini Morcaio Morchiaio Morcone Morellona di Grecia Moresca 1 Moresca 2 Morinello Murtiddara Nasitana frutto grosso Nebba

Toscany Toscany Toscany Toscany Toscany Apulia Sicily Calabria Calabria Marche Basilicata Basilicata Sardinia Sicily Sardinia Sicily Toscany Lazio Campania Toscany Toscany Toscany Toscany Toscany Apulia Marche Toscany Toscany Sicily Sicily Sicily Sicily Lazio Sicily Apulia Toscany Toscany Toscany Toscany Toscany Toscany Toscany Toscany Toscany Apulia Sicily Sicily Toscany Sicily Sicily Sicily

C1 # A1-134 A # 134 C1 # 287 C1 # 287 C1 # 288 A # 145 B # 194 A # 22 C1 # 321 C2 # 368 A # 12A A # 12B C2 # 378 A # 125 C2 # 379 B # 236 B # 203 A # 35 C2 # 360 A # 142 C2 #A4-135 C1 # A2-135 C1 # A3-135 A # 135 A # 60 A # 54 A # 136 C1 # 290 B # 211 A # 83A A # 83B A # 83c A # 36 B # 212 A # 131 C2 #A5-094 C1 # A3-094 C1 # A4-094 A # 94 B # A21 A # 137 C1 # 291 B # 203 C1 # 293 B # 178 A # 84A A # 84B B # 239 C2 # 385 B # 195 B # 259

251 252 253 254 255 256 257 258 259 260 261 262 263 264 265 266 267 268 269 270 271 272 273 274 275 276 277 278 279 280 281 282 283 284 285 286 287 288 289 290 291 292 293 294 295 296 297 298 299 300 301

Nebbia Nebbio di Chieti Nebbio di Pescara Negrera Nera di Cantinelle Nera di Gonnos Nera di Oliena Nera di Villacidro Nerba Nerba Catanese Noccioluta Nocellara del Belice 1 Nocellara del Belice 2 Nocellara etnea Nocellara etnea ovale Nocellara messinese Nocellara Nissena 1 Nocellara Nissena 2 Nociara Nolca Nostrale di Fiano Nostrale di Rigali Nostrana Nostrana Campana Nostrana di Brisighella 1 Nostrana di Brisighella 2 Nostrana di Brisighella 3 Nostrana di Brisighella 4 Ogliara Ogliarola Barese Ogliarola del Bradano 1 Ogliarola del Bradano 2 Ogliarola del Murge Ogliarola di Montalbano Ogliarola Garganica Ogliarola Messinese 1 Ogliarola Messinese 2 Ogliarola Salentina OgliarolaVulture Ogliastro grande Olianedda Oliastro Olieddu Oliva bianca Oliva da polpa Oliva grossa Oliva Rossa Oliva torsa Oliva vera Olivago Olivastra di Populonia

Marche Abruzzo Abruzzo Liguria Calabria Sardinia Sardinia Sardinia Sicily Sicily Molise Sicily Sicily Sicily Sicily Sicily Sicily Sicily Apulia Apulia Lazio Umbria Calabria Campania Emilia Romagna Emilia Romagna Emilia Romagna Emilia Romagna Campania Apulia Basilicata Basilicata Apulia Basilicata Apulia Sicily Sicily Apulia Basilicata Campania Sardinia Apulia Sardinia Campania Campania Emilia Romagna Apulia Campania Umbria Umbria Toscany

A # 105 A # 102 B # 176 A # 152 A # 23 A # 74 C1 # 312 A # 113 B # 258 C1 # 309 C2 # 371 A # 85 A # 85 A # 86 B # 183 A # 87 C1 # 351A C1 # 351B A # 61 A # 62 A # 37 A # 98 A # 24 B # 200 A # 148A A # 148B A # 148C A # A31 B # 245 A # 63 C2 # A1-013 A # 13 R P1F09 A # 14 B # 257 A # 88A A # 88B A # 64 A # 15 B # 246 C2 # 380 A # 112 C1 # 313 C2 # 361 C2 # 365 B # 199 C2 # 376 C2 # 362 C1 # 357 A # 155 C1 # 294

302 303 304 305 306 307 308 309 310 311 312 313 314 315 316 317 318 319 320 321 322 323 324 325 326 327 328 329 330 331 332 333 334 335 336 337 338 339 340 341 342 343 344 345 346 347 348 349 350 351 352

Olivastra Seggianese Olivastro d'Aprile Olivastro di Bucchianico Olivastro di Montenero Olivastro Dritto Olivastro Frentano Olivella appuntita Olivella di Cerchiara Olivella di Grottaminarda Olivetta Nera Olivo da olio Olivo da salare Olivo del mulino Olivo della Madonna Olivo delle Alpi Olivo di Casavecchia Olivo di Castiglione Olivo di San Lorenzo Olivoce Olivomensa Olivona Olivone di Viterbo OperaPia Orazio Orbetana Ornellaia Ortice Ortolana Ottobratica Ottobrina Paesana Bianca Paesana Nera Palma Palmarola Pampagliosa Paschixedda Pasola Passulunara Patrinostraru Pendolino 1 Pendolino 2 Pendolino 3 Pendolino 4 Pendolino 5 Pennulara Peppino Leo Peranzana Perciasacchi Pescarese Pesciatino Pezz'e quaddu

Toscany Molise Abruzzo Molise Molise Abruzzo Campania Calabria Campania Molise Campania Campania Toscany Toscany Liguria Toscany Sicily Toscany Abruzzo Basilicata Apulia Lazio Sicily Basilicata Marche Toscany Campania Campania Calabria Liguria Molise Molise Sardinia Basilicata Campania Sardinia Apulia Sicily Sicily Toscany Toscany Toscany Toscany Toscany Calabria Apulia Apulia Campania Abruzzo Toscany Sardinia

A # 173 C2 # 372 B # 218 C2 # 374 C2 # 373 B # 219 C1 # 327 C1 # 322 C2 # 363 C2 # 375 B # 247 B # 248 A # 140 A # 141 C1 # 342 C1 # 295 C1 # 310 C1 # 296 A#3 C1 # 267 R P1F03 C1 # 341 R P1F06 C2 # 392 A # 107 C1 # 297 B # 201 A # 161 A # 25 A # 48 B # 233 B # 234 C2 # 381 C1 # 268 A # 162 C1 # 314 A # 65 A # 143 I C2 # 386 C2 # A4-095 C1 # A3-095 C1 # A2-095 C1 # A1-095 A # 95 B # 225 A # 132 A # 66 B # 249 A#7 C1 # 298 C2 # 382

353 354 355 356 357 358 359 360 361 362 363 364 365 366 367 368 369 370 371 372 373 374 375 376 377 378 379 380 381 382 383 384 385 386 387 388 389 390 391 392 393 394 395 396 397 398 399 400 401 402 403

Piangente 1 Piangente 2 Piangente 3 Piantone di Falerone Piantone di Macerata Piantone di Mogliano Pidicuddara Pignola Pirunara 1 Pirunara 2 Pisciottana Pizz'e Carroga Pizzo di corvo 1 Pizzo di corvo 2 Pizzutella Pocciolo Posola Posolella Precoce Procanica Provenzale Puntella Punteruolo Puntino Racemo 1 Racemo 2 Racioppa Racioppa Racioppella Raggiola Raja Raja Sabina Rajo Rastellina Ravece Raza Razzaio Razzo 1 Razzo 2 Razzola Reale Regina Resciola di Venafro Riminino Ritonnella Rizza Rizzitella Roma Romanella Romanella Romanella Molisana

Toscany Toscany Toscany Marche Marche Marche Sicily Liguria Sicily Sicily Campania Sardinia Sicily Sicily Sicily Umbria Abruzzo Abruzzo Abruzzo Lazio Campania Abruzzo Toscany Toscany Apulia Apulia Basilicata Campania Campania Marche Umbria Lazio Umbria Umbria Campania Veneto Toscany Toscany Toscany Liguria Lazio Apulia Molise Lazio Campania Basilicata Campania Basilicata Basilicata Calabria Molise

C1 # A1-138 A # 138A A # 138B A # 106 C2 # 399 A # 108 A # 153 A # 150 C1 # 352A C1 # 352B A # 31 A # 75 C1 # 353A C1 # 353B B # 213 C1 # 356 B # 220 B # 221 A # 146 A # 38 B # 255 B # 177 C1 # 299 C1 # 300 C1 # 344 C1 # 344 C1 # 317 B # 250 A # 163 A # 109 A # 99 A # 149 A # 114 A # 172 C1 # 328 A # 156 C1 # 301 C2 # A1-169 A # 169 A # 49 A # 39 R P1F07 B # 189 A # 40 B # 251 A # 16 B # 252 C2 # 393 C2 # 394 A # 129 A # 167

404 405 406 407 408 409 410 411 412 413 414 415 416 417 418 419 420 421 422 423 424 425 426 427 428 429 430 431 432 433 434 435 436 437 438 439 440 441 442 443 444 445 446 447 448 449 450 451 452 453 454

Rosciola Rosciola Coltodino Rosciola di Rotello Rosciola di Venafro Rosciola laziale Rosino Rossellino Rossello Rossina Rotondella Rotondella di Foggia Rotondella di Sanza 1 Rotondella di Sanza 2 Rumignana Rustica Ruveia Salegna di Larino Salella Salicino Salvia Salviana Sammartinara Sammartinenga San Benedetto San Felice Acquasparta 1 San Felice Acquasparta 2 San Felice Acquasparta 3 San Felice Acquasparta 4 San Felice Acquasparta 5 San Francesco San Lazzero Sanginara Santa Caterina Santa Maria Sant'Agatese Sant'Agostino Santomauro Sargano Sargano Scarpetta Selvatico Semidana Sessana Simona Sinopolese Sirole Sivigliana da olio Spagnola di Missano Sperone di Gallo Spezzanese Spinoso

Marche Lazio Molise Molise Lazio Toscany Toscany Toscany Emilia Romagna Basilicata Apulia Campania Campania Molise Abruzzo Campania Molise Campania Toscany Lazio Lazio Sicily Basilicata Apulia Umbria Umbria Umbria Umbria Umbria Toscany Toscany Campania Toscany Campania Sicily Apulia Calabria Marche Marche Basilicata Emilia Romagna Sardinia Campania Apulia Calabria Lazio Sardinia Liguria Molise Calabria Basilicata

A # 110 B # 228 B # 186 B # 189 A # 41 C1 # 302 C1 # 303 C1 # 304 C1 # 335 C1 # 296 C2 # 377 C1 # 329A C1 # 329B B # 191 B # 222 C1 # 330 B # 235 C2 # 364 C1 # 306 A # 42 A # 43 B # 214 B # 224 B # 197 A # 100A A # 100B A # 100C A # 100D A # 100E A # 144 C1 # 305 C1 # 331 A # 96 B # 253 A # 89 A # 67 A # 160 A # 111 A # 165 C1 # 318 C1 # 336 A # 76 B # 256 A # 68 A # 27 A # 44 C1 # 345 C1 # 343 A # 175 C1 # 323 C2 # 395

455 456 457 458 459 460 461 462 463 464 465 466 467 468 469 470 471 472 473 474 475 476 477 478 479 480 481 482 483 484 485 486 487 488 489

Taggiasca Tenacella Tendellone Termite di Bitetto Terza grande Terza piccola Toccolana Tombarello Tonda di Alife Tonda di Cagliari Tonda di Filadelfia Tonda di Filogaso Tonda di Strongoli Tonda Dolce Tonda Dolce di Partanna Tonda iblea Tondina Tortiglione Toscanina Tunnulidda Turdunazza antimosca 1 Turdunazza antimosca 2 Uccellara Ugea di Savoia Ugliastride Vaddara Vallanella 1 Vallanella 2 Verdello Vigna della Corte Vocio Vraja Zaituna Zimbimbo Zinzifarica

Liguria Campania Umbria Apulia Sardinia Sardinia Abruzzo Calabria Campania Sardinia Calabria Calabria Calabria Calabria Sicily Sicily Calabria Abruzzo Apulia Sicily Sicily Sicily Basilicata Apulia Apulia Sicily Lazio Lazio Sicily Campania Umbria Calabria Sicily Apulia Calabria

A # 50 C1 # 332 A # 101 A # 69 A # 133 C1 # 316 A#8 B # 193 B # 254 A # 77 A # 28 A # 29 A # 116 B # 226 B # 237 A # 90 A # 30 A#9 A # 70 B # 215 C2 # 387A C2 # 387B A # 17 R P1F05 R P1F04 C1 # 354 A # 45A A # 45B A # 91 C1 # 333 B # 181 C2 # 396 A # 82A A # 123 A # 147

Table S2 Alleles detected at the SSR loci in the 489 olive trees analyzed. For each allele, the size (expressed in base pairs) and the frequency are reported. Locus

Alleles (bp) 1

2

3

4

5

GAPU59

208

212

214

218

222

Frequency

0.165 0.331 0.237 0.069 0.198

GAPU71A

210

Frequency

0.072 0.069 0.485 0.137 0.037 0.192

GAPU71B

121

Frequency

0.012 0.253 0.227 0.163 0.345

GAPU103A 136

212

124

150

214

126

157

218

130

159

221

6

7

8

9

224

228

250

259

10

11

12

220

232

243

0.001 0.002 0.005

144

170

184

197

203

Frequency

0.089 0.132 0.188 0.233 0.186 0.146

UDO01

140

Frequency

0.304 0.451 0.246

UDO03

135

Frequency

0.191 0.429 0.306 0.031 0.031 0.012

UDO12

156

Frequency

0.005 0.037 0.326 0.290 0.100 0.242

UDO28

143

Frequency

0.114 0.063 0.148 0.005 0.077 0.275

0.183 0.133 0.003

UDO39

108

205

Frequency

0.028 0.014 0.017 0.025 0.145 0.110

0.269 0.009 0.229

0.075 0.070

0.010

DCA09

162

188

204

210

Frequency

0.115 0.006 0.137 0.068 0.187 0.002

0.003 0.081 0.211

DCA18

163

181

Frequency

144

143

164

150

125

166

169

0.010 0.016

150

150

166

154

164

172

173

166

177

156

170

176

175

182

182

161

185

182

177

202

193

182

200

184

179

0.06 0.013 0.023 0.012 0.194 0.312

205

210

209

194

183

245

213

198

185

0.300 0.003 0.137

206

0.005 0.174

0.013

Table S3 Genotype profiles obtained from the combination of the eleven microsatellite markers on 489 olive varieties. The Italian olive varieties included in the core collection are indicated in bold. Varieties Abunara Agristigna Aitana Albatro Allora Amati Americano Annarea Arancino Arnasca 1 Arnasca 2 Arnasca 3 Arnasca 4 Arnasca 5 Arnasca 6 Ascolana dura Ascolana semitenera Ascolana tenera Augellina Aurina Barilari Bianchella Bianchera Biancolella Biancolilla Borgese Borgiona Bosana Bottone di Gallo 1 Bottone di Gallo 2 Bottoni di Gallo Brandofino Buga

GAPU59 GAPU71A GAPU71B GAPU103A UDO001 UDO003 UDO012 UDO028 UDO039 DCA09 DCA18 208-212 210-210 124-144 159-170 144-144 143-143 177-177 182-182 108-108 162-162 179-181 208-208 210-210 124-144 170-184 144-144 143-143 166-177 143-205 200-200 198-206 179-181 208-212 210-210 124-144 150-184 144-144 143-143 166-193 182-182 108-108 176-176 179-179 208-222 214-214 124-144 159-170 144-144 150-150 177-177 143-182 205-205 176-206 179-181 212-212 214-214 126-144 159-159 150-150 143-143 164-164 154-182 164-205 206-206 181-185 214-218 214-224 124-144 157-159 144-144 150-150 166-177 150-161 213-213 182-198 177-179 212-218 218-224 126-126 159-170 144-144 150-150 177-193 182-182 205-205 162-198 179-181 212-222 214-224 124-144 184-184 144-144 150-150 166-193 182-210 205-205 182-182 179-181 212-218 214-214 124-130 184-184 140-140 150-150 156-193 154-205 220-243 172-194 179-185 212-212 224-224 126-144 159-184 140-140 150-150 166-193 154-205 213-232 182-206 181-185 212-212 224-224 126-144 159-184 140-140 150-150 166-193 154-205 213-232 182-206 181-185 212-212 224-224 126-144 159-184 140-140 150-150 166-193 154-205 213-232 182-206 181-185 208-212 214-259 124-144 157-157 144-144 150-150 166-193 154-154 213-213 182-206 181-185 212-212 224-224 126-144 159-184 140-140 150-150 166-193 154-205 213-232 182-206 181-185 212-212 214-218 130-144 170-184 140-140 135-135 164-177 161-205 213-232 162-176 177-179 214-222 214-224 124-130 136-157 140-140 143-143 166-166 182-210 200-200 198-206 177-181 214-222 214-224 124-130 136-157 140-140 135-135 166-166 182-210 200-232 198-206 177-181 214-222 214-224 126-144 136-184 140-140 143-143 166-166 150-161 200-200 198-206 177-179 208-208 210-210 124-126 157-203 140-140 143-143 164-166 150-150 205-205 194-198 177-181 222-222 210-214 124-144 159-170 144-144 150-150 177-177 154-205 213-213 162-206 173-175 212-212 214-214 124-130 159-184 144-144 150-150 166-193 154-182 205-205 172-198 177-179 212-222 218-224 130-144 157-157 140-140 135-135 177-182 182-210 185-220 176-206 181-185 214-214 214-224 130-130 136-157 140-140 150-150 166-193 182-205 213-232 182-182 177-185 212-214 214-224 124-144 159-170 144-150 150-150 164-177 182-210 185-185 182-206 181-185 212-212 210-214 121-130 136-150 144-144 150-150 166-193 143-205 108-108 172-198 177-179 212-222 214-214 130-144 150-184 144-144 143-143 166-193 143-205 200-220 162-198 179-179 214-222 214-224 130-144 157-157 140-140 143-143 166-193 150-210 200-200 162-198 177-181 214-222 214-218 130-144 170-184 144-150 135-135 177-193 161-205 205-205 194-198 181-185 212 -222 214-259 124-126 150-184 144-144 150-150 177-193 154-182 220-220 162-172 179-185 212 -222 214-259 124-126 150-184 144-144 150-150 177-193 154-182 220-220 162-182 179-185 212-212 210-214 124-144 170-170 144-144 150-150 177-193 154-205 213-213 172-206 173-177 208-212 210-214 126-144 136-159 144-144 150-150 166-193 143-143 213-232 176-206 179-179 214-214 218-218 126-144 136-157 140-140 135-135 166-193 182-205 200-200 182-198 179-185

Buscionetto 1 Buscionetto 2 Buscionetto 3 Cacaredda Cacaridduni Cacazzara Caiazzana Calatina Cammarotana Canino 1 Canino 2 Capolga Caprina di Casalanguida Caprina Vastese Carbona Carboncella Carboncella Campana Carboncella Pianacce A Carboncella Pianacce B Carbonchio Carbuncion Carbuncion di Carpinenga Cariasina Cariola Carmelitana Carolea Cefaly Carolea Cetraro Carolea Mirto Carolea Rende Carolea Rossi Carpellese Carpinella Carpinetana Casaliva Castiglionese Castricianella Rapparina Cavalieri

212 -222 212-212 212-212 208-212 208-212 208-212 218-222 212-222 214-222 212 -222 212 -222 214-214 208-212 208-212 214-214 208-212 214-218 208-212 208-212 212-218 218-222 214-218 214-214 218-222 208-208 212-222 212-222 212-222 212-222 212-222 214-222 212-218 212-222 208-212 208-212 208-222 212-222

210-214 210-214 210-214 210-210 210-210 214-214 214-214 210-214 212-218 214-221 214-221 214-224 214-214 214-214 214-224 214-224 218-218 214-214 214-214 214-214 218-218 214-214 214-218 212-212 214-224 214-214 214-214 214-214 214-214 214-214 214-224 214-214 214-214 214-224 210-214 210-214 210-210

124-130 124-126 124-126 124-144 124-144 124-126 126-126 124-126 124-144 126-144 126-144 124-144 124-124 124-130 126-130 124-144 124-126 124-144 124-130 124-124 126-144 126-144 130-144 130-144 124-126 121-130 130-144 121-130 121-130 130-144 124-144 126-144 126-144 124-144 126-126 124-130 124-144

159-184 136-150 136-150 159-184 150-184 157-184 170-184 136-136 159-170 159-159 159-159 136-150 150-150 150-150 150-157 170-184 150-157 170-184 157-157 150-170 159-159 157-159 136-184 184-184 136-159 136-170 136-170 136-170 136-184 136-170 157-159 150-159 184-184 159-170 159-184 159-170 159-159

144-144 140-140 140-140 144-144 144-144 144-144 140-140 144-144 140-140 150-150 150-150 144-144 144-144 144-144 150-150 144-144 144-150 144-144 144-144 140-140 150-150 140-140 144-150 144-144 144-144 144-144 144-144 144-144 144-144 144-144 144-144 150-150 144-144 144-144 144-144 144-144 144-144

150-150 150-150 150-150 143-143 143-143 150-150 135-135 150-150 143-143 135-135 135-135 150-150 150-182 150-202 143-143 150-150 150-150 135-182 166-166 150-150 150-150 135-135 143-143 150-150 143-143 143-143 143-143 143-143 143-143 143-143 143-143 150-150 150-150 143-143 150-150 150-150 150-150

166-166 166-193 166-193 166-193 166-193 166-193 164-177 177-177 164-177 166-182 166-182 166-193 166-193 177-193 166-193 166-193 177-177 166-166 177-193 177-193 166-182 177-182 166-193 166-193 166-193 166-193 166-193 166-193 166-193 166-193 166-177 166-177 177-193 177-182 166-193 177-177 177-193

154-154 154-154 154-154 182-182 182-182 182-182 143-143 143-182 143-143 143-154 143-154 154-205 154-205 154-205 143-210 182-210 150-182 143-210 154-205 154-205 182-205 182-182 150-161 143-154 182-182 154-205 143-205 154-205 154-205 143-205 161-210 156-205 154-210 182-205 154-210 154-205 154-154

108-213 213-213 213-213 205-205 108-108 185-185 200-213 205-205 213-213 213-232 213-232 220-232 213-213 213-213 185-185 213-213 200-200 205-205 213-213 213-213 125-125 125-200 205-205 205-205 185-185 200-200 200-200 200-200 125-213 200-200 200-200 205-205 205-205 205-205 205-232 170-213 108-108

176-198 176-198 176-198 176-206 162-176 176-176 198-198 182-182 194-206 172-182 172-182 176-198 162-204 162-204 176-198 162-194 182-206 162-194 162-194 198-210 176-182 172-198 162-172 162-198 198-198 162-198 162-198 162-198 162-198 162-198 172-198 182-182 166-206 172-194 198-206 198-206 162-162

179-181 181-181 181-181 181-181 169-181 179-179 179-181 175-179 179-181 173-179 173-179 181-185 177-177 177-177 179-181 177-181 181-185 177-181 177-181 181-185 177-181 177-181 179-181 181-185 179-181 179-185 179-185 179-185 179-181 179-185 181-185 181-181 177-181 179-181 163-181 179-179 177-185

Cazzarella Cazzinicchio Cellacchia Cellina di Nardo' Cellina di Rotello Cerasa di Montenero Cerasuola Ciciarello Cicinella Ciliegino Cima di Melfi Cima di Mola Colombina Coratina 1 Coratina 2 Coratina 3 Cornacciola Corneglia Cornia Corniola Corniolo Cornula Coroncina Correggiolo Correggiolo di Montegridolfo Correggiolo di Pallese Corsicana da mensa Corsicana da olio Craputea Crastu Crognolo CSM CSS-02-Minerva Cucca 1 Cucca 2 Curatora Demoniata

212-212 212-212 212-218 208-222 222-222 212-222 208-212 212-212 214-218 208-208 212-222 212-212 212-222 212-212 212-212 212-212 212-222 218-222 214-218 208-212 214-222 208-212 212-222 208-212 208-212 208-212 214-222 212-222 214-214 208-208 214-222 208-212 208-212 208-212 212-212 218-222 212-212

214-214 210-214 218-224 214-228 210-214 214-214 210-214 212-212 212-218 214-218 210-224 210-214 218-224 210-224 210-224 210-224 214-214 214-224 218-218 212-212 214-214 210-214 212-214 214-224 214-224 214-224 212-221 214-224 218-218 214-214 218-224 214-224 210-214 214-214 214-214 218-218 210-214

121-121 124-126 124-144 124-126 124-126 126-144 124-144 124-124 124-130 126-144 124-144 124-144 144-144 124-144 124-144 124-144 126-144 126-126 130-144 124-144 130-144 124-144 130-144 124-144 124-144 124-144 126-144 126-144 126-126 124-144 124-144 124-124 130-144 124-144 124-144 126-126 124-130

170-170 159-159 150-184 170-170 150-184 170-184 150-184 159-170 170-184 170-184 170-170 159-159 184-203 136-170 136-170 136-170 150-159 136-170 184-197 170-184 150-157 157-157 170-184 159-170 159-170 159-170 157-159 159-184 157-159 157-170 157-170 159-159 184-184 170-170 170-170 170-184 159-159

140-140 140-140 144-144 144-144 140-140 144-144 144-144 144-144 140-140 144-144 144-144 144-144 150-150 140-140 140-140 140-140 150-150 140-140 140-140 144-144 140-140 140-140 150-150 144-144 144-144 144-144 150-150 144-150 150-150 144-144 150-150 144-144 144-144 144-144 144-144 144-150 140-140

143-143 143-143 135-135 150-150 202-202 143-143 150-150 143-143 143-143 202-202 150-150 143-143 143-143 135-166 135-166 135-166 143-143 143-143 143-143 143-143 135-143 150-150 135-135 143-143 143-143 143-143 143-143 143-143 143-143 150-150 150-150 150-150 150-150 150-150 150-182 135-135 150-150

166-193 166-193 166-166 166-166 177-193 177-193 166-166 177-193 166-193 156-166 166-193 177-193 166-193 166-193 166-193 166-193 166-177 166-177 164-177 177-193 177-193 177-193 177-193 177-182 177-182 177-182 166-166 177-182 177-193 177-177 177-182 177-177 166-166 177-193 177-177 166-166 177-193

182-182 143-205 143-143 182-205 154-210 143-182 154-154 143-143 154-161 156-205 154-210 150-182 182-182 143-210 143-210 143-210 156-205 154-182 143-154 143-205 161-205 182-205 161-182 182-205 182-205 182-205 150-161 182-210 182-182 182-182 182-210 182-210 154-205 182-182 182-210 143-143 182-182

213-213 108-205 205-205 108-213 170-213 205-205 213-213 205-205 205-205 220-232 170-213 170-170 213-213 170-170 170-170 170-170 205-205 213-220 164-213 200-200 205-205 220-243 200-200 205-205 205-205 205-205 213-213 185-185 185-185 185-185 164-164 213-213 205-205 213-213 205-205 185-185 108-185

162-172 182-198 182-198 176-198 194-206 176-206 176-206 198-198 172-198 172-194 194-194 162-162 172-206 172-194 172-194 172-194 176-182 172-198 198-198 172-182 182-206 162-182 172-194 172-194 172-194 172-194 176-182 182-198 172-194 182-198 176-198 162-206 162-206 162-198 162-198 198-206 176-206

169-179 181-181 181-181 177-177 177-181 173-179 177-181 177-177 185-185 179-185 181-181 173-177 179-179 181-181 181-181 181-181 181-185 181-181 181-181 177-179 179-181 179-185 181-185 179-181 179-181 179-181 179-181 181-185 179-185 179-181 179-181 177-177 177-179 179-181 179-181 179-185 181-185

Dolce Agogia Dolce di Andria Dolce di Cassano Dolce di Cerchiara Dolce di Rossano Don Carlo Donna Francesca Dritta Dritta Loreto Emilia Erbano 1 Erbano 2 Faresana Faresana Fasolina Fasolona Favarol Fecciaro Feglina Femminella di Torraca Filare Fosco Frangivento Frantoio 1 Frantoio 10 Frantoio 2 Frantoio 3 Frantoio 4 Frantoio 5 Frantoio 6 Frantoio 7 Frantoio 8 Frantoio 9 Frantoio di Montegridolfo Frantoio di Villa Verrucchio Frantoio FC Fs17

212-218 208-218 208-208 212-222 212-212 212-212 208-222 208-212 214-222 208-212 212-212 212-222 212-212 208-208 214-222 214-214 208-208 214-222 212-212 218-222 212-222 208-212 208-208 208-212 208-212 208-212 208-212 208-212 208-212 208-212 208-212 208-212 208-212 208-212 208-212 208-212 214-214

218-218 214-214 214-224 212-212 214-214 221-221 210-224 214-214 214-224 214-214 210-212 214-250 214-214 214-214 214-214 214-214 214-214 214-221 212-212 214-218 214-214 214-224 214-214 214-224 214-224 214-224 214-224 214-224 214-224 214-224 214-224 214-224 214-224 214-224 214-224 214-224 214-224

124-124 124-144 130-130 130-144 130-144 124-144 124-144 124-130 144-144 124-144 124-144 124-144 124-126 124-126 126-144 130-130 126-144 124-130 126-144 130-144 124-130 124-144 124-130 124-144 124-144 124-144 124-144 124-144 124-144 124-144 124-144 124-144 124-144 124-144 124-144 124-144 124-124

184-197 150-157 150-157 136-170 150-159 136-157 136-159 159-170 159-159 159-170 159-159 159-170 159-184 159-184 150-159 159-184 150-157 159-159 170-203 157-159 150-150 159-170 159-170 159-170 159-170 159-170 159-170 159-170 159-170 159-170 159-170 159-184 159-184 159-170 159-170 159-170 184-197

140-140 144-144 144-144 150-150 144-144 144-144 144-144 140-140 144-144 144-144 140-140 140-140 150-150 140-140 150-150 150-150 150-150 140-140 150-150 140-140 144-144 144-144 144-144 144-144 144-144 144-144 144-144 144-144 144-144 144-144 144-144 144-144 144-144 144-144 144-144 144-144 140-140

143-143 143-143 143-143 143-143 143-143 150-150 150-150 143-143 150-150 150-150 150-150 150-150 143-143 143-143 150-150 150-150 143-143 143-143 135-135 143-143 150-150 150-150 135-166 143-143 143-143 143-143 143-143 143-143 143-143 143-143 143-143 143-143 143-143 143-143 143-143 143-143 143-143

177-193 177-182 177-177 166-193 177-177 166-193 177-193 166-193 177-193 156-193 177-193 177-193 166-177 166-177 177-177 177-193 166-177 166-177 177-193 164-177 177-193 177-177 177-177 177-182 177-182 177-182 177-182 177-182 177-182 177-182 177-182 177-182 177-182 177-182 177-182 177-182 166-177

143-143 143-205 182-205 143-205 154-182 154-205 154-205 182-182 182-210 182-182 182-182 143-182 150-150 143-161 154-205 154-154 154-210 143-182 143-205 143-143 154-205 182-210 154-205 182-205 182-205 182-205 182-205 182-205 182-205 182-205 182-205 182-205 182-205 182-205 182-205 182-205 150-205

185-213 108-170 108-170 205-205 205-205 125-200 170-205 220-220 185-220 213-213 108-108 213-213 205-205 205-205 205-205 205-205 185-185 205-205 164-213 164-213 213-213 205-205 205-232 205-205 205-205 205-205 205-205 205-205 205-205 205-205 205-205 205-205 205-205 205-205 205-205 205-205 185-220

172-182 172-194 162-198 162-198 172-182 198-206 182-198 172-176 172-206 172-206 172-182 172-182 182-198 182-198 198-206 198-206 182-206 176-198 162-176 198-206 184-206 172-194 162-176 172-194 172-194 172-194 172-194 172-194 172-194 172-194 172-194 172-194 172-194 172-194 172-194 172-194 206-206

179-181 177-177 181-181 179-181 177-179 177-181 179-179 169-179 181-185 177-179 173-177 173-177 179-181 179-181 179-179 179-181 177-177 179-181 181-181 181-181 179-185 179-181 179-185 179-181 179-181 179-181 179-181 179-181 179-181 179-181 179-181 179-181 179-181 179-181 179-181 179-181 179-181

Gaggiolo Gentile dell'Aquila Gentile di Chieti Gentile di Larino 1 Gentile di Larino 5 Gentile nera di Colletorto Ghiacciolo Ghiacciora Ghiandaro Ghiannara Giardino Giarfara Giarraffa 1 Giarraffa 2 Ginestrino Giusta Gnagnaro Grappolo Grappuda Gremignolo Gremignolo di Bolgheri Grignan Grossa di Cassano Grossa di Gerace Grossa di Spagna Grossa di Venafro Grossale I 77 Iacona 1 Iacona 2 Imperiale Intosso Itrana Larcianese Lastrino Laurina Lavagnina

214-222 212-212 208-212 208-212 208-212 208-212 214-214 212-212 208-212 212-218 214-222 212-212 212-212 212-212 208-212 214-222 212-214 208-218 212-212 208-212 222-222 212-212 208-218 212-212 208-208 212-222 214-214 218-222 208-212 208-212 208-212 212-222 214-218 208-218 212-218 214-214 212-212

214-224 214-214 210-214 214-214 214-214 214-214 214-224 214-214 214-214 214-224 218-218 214-214 210-214 214-214 214-224 214-214 210-214 210-218 218-218 214-224 214-224 212-221 210-214 212-212 210-214 214-214 214-214 218-218 214-214 214-214 210-250 210-214 214-224 212-221 210-210 214-224 210-214

126-144 144-144 124-130 124-130 124-130 124-126 126-144 124-144 124-144 124-144 126-144 144-144 124-144 144-144 144-144 126-144 124-144 124-144 130-144 124-144 144-144 124-130 124-144 126-144 124-144 124-130 126-144 124-130 124-144 124-144 130-144 124-130 130-144 124-144 124-130 124-144 130-144

159-170 150-184 170-170 150-184 150-184 136-170 157-184 136-157 159-170 170-184 170-184 157-184 136-159 150-184 159-184 170-184 170-170 170-184 184-203 170-184 159-184 157-157 136-159 150-170 136-184 150-150 157-159 150-157 159-184 159-184 159-184 159-184 159-170 159-170 184-203 157-157 136-170

150-150 140-140 140-140 140-140 140-140 140-140 140-140 150-150 144-144 144-144 150-150 140-140 144-144 140-140 144-144 144-144 144-144 150-150 140-140 144-144 144-144 140-140 140-140 140-150 144-144 144-144 144-144 140-140 144-144 144-144 144-144 140-140 150-150 150-150 144-144 144-144 144-144

143-143 182-182 143-143 150-150 150-150 135-135 135-135 135-135 150-150 150-150 150-150 150-150 150-150 150-150 150-150 143-143 150-182 150-150 150-150 150-150 150-150 150-150 143-143 143-143 143-143 150-150 143-150 135-135 150-150 150-150 150-150 166-166 143-143 143-143 135-135 143-143 150-182

177-182 166-193 166-193 166-193 166-193 177-193 166-166 166-177 177-193 177-177 166-193 166-193 166-166 166-193 177-193 177-177 166-166 156-182 166-193 156-193 156-193 166-166 166-177 166-177 166-166 177-193 166-166 166-177 166-193 166-193 166-193 166-166 166-177 166-177 156-166 177-193 166-193

182-205 182-205 154-205 182-182 182-182 154-205 150-161 150-210 154-205 182-210 182-182 182-182 143-182 182-182 182-205 182-182 154-182 182-182 161-205 182-205 182-205 182-210 182-182 143-182 143-245 154-205 143-143 154-205 182-182 182-182 143-143 154-205 150-161 161-205 156-161 154-205 143-245

185-185 213-213 213-213 213-213 213-213 213-213 185-232 205-205 205-205 205-205 213-213 108-108 213-213 108-108 220-220 205-205 220-243 220-220 213-232 205-205 205-205 213-220 205-232 200-200 108-108 213-213 205-232 205-205 108-205 108-205 213-232 205-205 220-220 200-200 220-232 200-200 213-232

182-206 198-210 172-172 172-194 172-194 198-210 198-206 162-176 166-182 176-206 182-206 176-182 162-198 172-182 162-198 162-198 162-206 182-206 172-182 194-206 194-206 194-198 162-182 182-198 162-172 162-204 194-198 198-206 162-162 162-162 182-198 162-210 182-198 182-206 194-198 172-172 172-176

179-179 175-181 169-179 169-179 169-179 177-179 177-179 179-181 163-177 179-185 181-181 179-179 181-185 177-181 181-181 177-179 175-175 181-181 177-181 177-179 177-179 177-185 179-179 179-181 177-177 177-177 179-181 179-179 179-181 179-181 177-185 179-181 179-181 181-181 179-179 177-185 179-181

Lazzera Reale Lea Leccino 1 Leccino 10 Leccino 2 Leccino 3 Leccino 4 Leccino 5 Leccino 6 Leccino 7 Leccino 8 Leccino 9 Leccino Compatto Leccino Dwarf Leccino Minerva 1 Leccino Minerva 2 Leccino Pisa 4 Leccino Pisa 9 Leccio del Corno 1 Leccio del Corno 2 Leccio Marammano 1 Leccio Maremmano 2 Leccione Lezze Lumiaru Mafra Mafra di Cerchiara Maglianese Maiatica di Ferrantina 1 Maiatica di Ferrantina 2 Majorca Mandanici Manna Mantonica Marchiaio Marina Marinese

212-214 214-214 212-212 212-212 212-222 212-222 212-222 212-222 212-222 212-222 212-222 212-222 212-222 208-212 208-212 212*-212 208-212 208-212 208-212 208-212 208-212 208-212 208-212 208-218 208-212 212-212 208-222 212-212 208-208 208-208 214-214 208-212 214-214 208-222 208-208 212-222 212-214

214-214 218-224 214-214 214-224 214-224 214-214 214-214 214-224 214-224 214-224 214-224 214-224 214-224 214-224 214-224 214-224 214-224 214-224 214-224 214-214 214-224 214-224 214-224 210-210 214-214 212-212 212-221 214-224 214-214 214-214 214-224 210-214 214-224 214-214 210-210 214-214 212-224

124-126 130-144 124-130 124-130 124-144 124-130 124-130 124-130 126-144 126-144 126-144 126-144 124-144 124-144 124-144 144-144 124-144 124-144 124-130 144-144 124-144 124-144 124-124 124-144 124-144 126-144 126-144 126-144 124-144 124-144 126-144 124-130 126-144 124-126 124-144 126-126 124-144

157-157 159-159 150-150 150-150 150-150 150-150 150-150 150-150 184-203 184-203 184-203 184-203 150-150 170-184 170-184 159-203 170-184 170-184 150-150 159-184 170-184 170-184 159-159 136-159 159-184 136-159 136-159 157-157 170-184 170-184 170-184 159-170 170-184 157-159 150-157 184-197 136-136

140-140 140-140 144-144 144-144 144-144 144-144 144-144 144-144 144-144 144-144 144-144 144-144 144-144 144-144 144-144 144-144 144-144 144-144 144-144 144-144 144-144 144-144 144-144 140-140 144-144 144-144 144-144 140-140 144-144 144-144 150-150 144-144 150-150 144-144 150-150 144-144 144-144

150-150 143-150 143-143 143-143 143-143 143-143 143-143 143-150 150-150 150-150 150-150 150-150 143-143 150-150 150-150 150-150 150-150 150-150 166-166 150-150 150-150 150-150 150-150 135-135 143-143 143-143 143-143 143-143 143-143 143-143 135-135 150-150 135-135 150-150 166-166 135-135 135-143

156-166 166-193 166-193 166-193 177-193 166-193 166-193 177-193 166-193 166-193 166-193 166-193 177-193 166-193 166-193 166-193 166-193 166-193 177-177 166-193 166-193 166-193 177-177 177-182 166-193 166-182 166-177 166-182 166-193 166-193 166-193 177-193 166-193 177-177 164-182 166-193 166-166

154-182 154-210 154-205 154-205 154-205 154-205 154-205 154-154 182-210 182-210 182-210 182-210 154-205 182-210 182-210 182-210 182-210 182-210 154-205 182-210 182-182 182-210 182-210 143-161 143-182 143-143 143-205 154-210 182-182 182-182 150-150 143-210 150-150 182-182 182-182 154-154 143-210

220-232 205-232 213-213 213-213 213-220 213-213 213-213 213-213 205-205 205-205 205-205 205-205 213-213 205-205 205-205 205-205 205-205 205-205 205-205 205-205 205-205 205-205 205-232 164-185 205-205 213-213 213-213 185-232 205-205 205-205 213-220 164-205 213-220 108-205 125-125 213-232 205-213

162-206 198-206 162-206 162-206 162-206 162-206 162-206 162-206 162-206 162-206 162-206 162-206 162-206 162-206 162-206 162-206 162-206 162-206 182-206 182-206 198-206 198-206 172-206 172-194 176-206 162-198 162-182 198-206 198-206 198-206 162-172 162-182 162-172 172-182 194-194 162-198 162-194

179-181 179-181 177-177 177-177 177-177 177-177 177-177 177-177 177-177 177-177 177-177 177-177 177-177 177-177 177-177 177-177 177-177 177-177 181-181 181-181 177-181 177-181 179-185 181-181 177-181 179-179 179-185 177-181 177-179 177-179 181-185 179-181 181-185 177-181 179-181 179-181 179-179

Marzio Maurino 1 Maurino 2 Maurino 3 Maurino 4 Mele Mignola Mignolo Mignolo Cerretano Minna di Vacca Minuta 1 Minuta 2 Minuta3 Minutella Monaca Mora Moraiolo 1 Moraiolo 2 Moraiolo 3 Moraiolo 4 Moraiolo 5 Moraiolo T. Corsini Morcaio Morchiaio Morcone Morellona di Grecia Moresca 1 Moresca 2 Morinello Murtiddara Nasitana frutto grosso Nebba Nebbia Nebbio di Chieti Nebbio di Pescara Negrera Nera di Cantinelle

208-218 208-208 208-212 208-212 208-212 208-208 214-218 208-218 208-218 208-212 208-208 208-212 212-212 214-214 208-212 208-212 212-222 212-222 212-222 212-222 212-222 212-222 212-218 218-218 208-222 212- 222 208-212 208-212 212-214 208-208 208-212 208-208 208-212 212- 222 208-208 214-214 212-222

210-218 214-224 214-224 214-224 214-224 214-214 214-221 210-218 214-224 214-214 210-214 210-214 214-221 214-224 210-214 210-224 214-214 214-214 214-214 214-214 214-214 214-214 210-210 212-212 210-214 210-214 214-214 214-214 218-218 210-214 214-214 214-214 210-214 214-214 210-214 214-224 212-212

130-130 144-144 144-144 144-144 130-144 124-144 126-144 124-144 124-126 124-130 126-126 124-144 124-124 124-130 124-144 124-126 124-130 124-124 124-130 124-124 124-124 124-130 126-144 124-144 124-126 124-126 124-144 124-144 130-144 144-144 124-130 144-144 124-130 124-130 124-130 126-144 126-144

159-159 159-184 184-184 159-184 159-184 150-157 157-157 159-170 150-159 159-184 157-157 159-170 159-159 136-150 157-159 170-184 150-150 150-150 150-157 150-150 150-157 150-157 150-150 150-157 157-157 136-136 159-184 159-184 150-157 159-159 159-170 150-184 159-184 159-159 170-170 170-184 136-150

150-150 144-144 144-144 144-144 144-144 144-144 144-144 150-150 140-140 140-140 144-144 144-144 150-150 144-150 144-144 144-144 144-144 144-144 144-144 144-144 144-144 144-144 150-150 140-140 144-144 140-140 144-144 144-144 140-140 144-144 144-144 140-140 140-140 144-144 140-140 150-150 144-144

150-150 150-150 150-150 150-150 150-150 143-143 143-143 150-150 143-143 150-150 143-166 143-166 150-150 135-135 150-150 143-143 182-182 182-182 182-182 182-182 166-182 166-182 143-143 143-143 135-166 166-166 166-202 166-202 150-150 150-150 143-143 150-150 135-135 135-135 166-166 143-143 143-143

166-182 177-182 177-193 177-182 177-193 166-177 177-193 156-177 177-177 166-193 177-193 177-177 177-193 166-166 177-193 166-193 177-193 177-193 177-193 177-193 177-193 177-193 156-193 156-177 177-193 166-166 166-193 166-193 177-193 177-193 166-193 166-193 177-177 177-193 166-193 166-193 166-193

182-205 182-210 182-210 182-210 182-210 150-182 154-205 182-205 182-205 182-182 182-182 154-205 154-245 150-161 154-182 143-210 154-205 154-154 154-205 154-205 154-205 154-205 154-182 150-182 182-205 182-182 143-182 143-182 154-182 182-182 210-210 182-182 154-210 154-205 154-210 182-182 143-143

200-220 220-220 220-220 220-220 220-220 108-170 200-200 200-200 164-164 108-108 170-213 170-170 164-213 205-205 164-205 108-164 213-213 213-213 213-213 213-213 213-213 213-220 205-205 108-108 220-243 108-205 108-213 108-213 200-200 205-205 205-205 205-205 205-205 213-213 213-232 200-200 200-200

162-198 162-198 162-198 162-198 162-198 162-182 172-172 182-206 162-172 176-206 176-198 162-198 176-206 162-198 176-206 162-182 184-206 184-206 184-206 184-206 184-206 184-206 182-206 194-194 176-176 182-198 182-198 182-198 182-206 182-198 162-198 176-206 172-206 172-210 172-182 162-198 162-198

181-181 181-181 181-181 181-181 181-181 177-185 177-185 181-181 175-181 181-185 179-179 179-179 181-181 179-181 181-181 179-181 179-185 179-185 179-185 179-185 179-185 179-185 185-185 177-181 177-181 179-185 177-179 177-179 179-181 177-181 177-185 181-185 177-181 175-179 169-179 179-179 177-179

Nera di Gonnos Nera di Oliena Nera di Villacidro Nerba Nerba Catanese Noccioluta Nocellara del Belice 1 Nocellara del Belice 2 Nocellara etnea Nocellara etnea ovale Nocellara messinese Nocellara Nissena 1 Nocellara Nissena 2 Nociara Nolca Nostrale di Fiano Nostrale di Rigali Nostrana Nostrana Campana Nostrana di Brisighella 1 Nostrana di Brisighella 2 Nostrana di Brisighella 3 Nostrana di Brisighella 4 Ogliara Ogliarola Barese Ogliarola del Bradano 1 Ogliarola del Bradano 2 Ogliarola del Murge Ogliarola di Montalbano Ogliarola Garganica Ogliarola Messinese 1 Ogliarola Messinese 2 Ogliarola Salentina OgliarolaVulture Ogliastro grande Olianedda Oliastro

212-212 214-222 214-222 208-212 208-212 208-212 212-212 212-212 212-212 212-212 208-212 208-212 208-212 212-222 208-208 212-222 214-214 212-222 218-222 214-214 212-212 214-214 212-212 214-222 208-212 212-212 208-212 208-212 208-212 208-212 208-212 208-212 212-212 212-218 214-218 214-222 208-218

214-221 212-218 214-221 214-214 214-214 214-214 214-214 214-214 210-210 210-210 214-214 210-214 210-214 210-214 214-214 214-224 218-224 212-212 218-218 214-224 214-214 214-224 214-214 214-214 214-224 210-214 214-224 214-224 214-224 214-224 214-214 214-214 210-214 214-214 218-218 214-221 210-210

126-144 126-144 126-144 124-144 124-144 124-126 144-144 144-144 124-144 124-144 124-144 124-144 124-144 124-126 124-144 126-144 124-130 124-130 126-144 126-144 124-130 126-144 124-126 124-126 124-144 124-144 124-144 124-144 124-144 124-144 124-144 124-144 124-144 124-124 126-126 126-144 126-144

170-203 157-159 157-159 159-184 159-184 136-170 150-184 150-184 159-184 159-184 159-184 136-159 136-159 170-170 150-157 159-170 184-197 170-184 184-197 136-184 136-157 136-184 136-157 150-157 159-170 159-159 159-170 159-184 159-170 159-170 159-184 159-184 159-159 150-184 159-159 157-159 136-159

150-150 144-150 144-150 144-144 144-144 140-140 140-140 140-140 144-144 144-144 144-144 144-144 144-144 140-140 144-144 144-144 144-144 140-150 140-140 144-144 144-144 144-144 144-144 140-140 144-144 144-144 144-144 144-144 144-144 144-144 144-144 144-144 144-144 144-150 144-144 144-144 140-140

135-135 143-143 143-143 150-150 150-150 182-182 166-166 166-166 150-150 143-143 150-150 150-150 150-150 143-143 143-143 143-143 135-143 143-150 135-150 135-135 135-135 135-135 135-135 143-143 143-143 150-150 143-143 150-150 143-143 143-143 166-182 166-182 143-143 143-143 150-150 143-143 143-143

166-193 164-177 164-177 166-193 166-193 177-193 166-193 166-193 166-193 166-193 166-193 166-193 166-193 166-193 166-177 177-182 166-182 177-177 166-182 166-166 166-193 166-166 166-193 164-193 177-182 177-193 177-182 177-177 177-182 177-182 166-193 166-193 177-193 166-193 164-164 166-177 166-177

143-143 150-161 150-161 154-205 154-182 154-205 182-182 182-182 143-182 143-182 143-143 154-154 154-154 182-182 150-182 182-210 150-161 143-143 143-210 150-161 182-210 150-161 182-210 161-161 182-205 154-205 182-205 182-205 182-205 182-205 182-182 182-182 150-182 182-182 143-143 150-161 143-161

213-220 213-213 213-213 213-213 213-213 213-213 108-205 108-205 220-220 220-220 213-232 205-205 205-205 164-232 108-170 185-185 205-220 205-205 164-213 213-213 205-220 213-213 205-220 213-232 205-205 205-205 205-205 205-205 205-205 205-205 108-205 108-205 170-170 213-213 200-220 213-213 164-185

162-172 176-182 182-198 176-206 162-172 198-210 162-172 162-172 172-182 198-206 162-176 176-198 176-198 162-162 162-182 182-206 162-172 172-206 194-198 198-198 198-198 198-206 198-198 172-198 172-194 172-172 172-194 172-198 172-194 172-194 162-162 182-198 162-162 172-182 182-206 176-182 172-194

185-185 179-181 179-181 181-185 177-185 177-181 177-181 177-181 181-181 181-185 179-179 179-179 179-179 179-185 177-185 179-181 179-181 177-179 181-181 177-179 177-179 179-185 177-179 179-185 179-181 177-185 179-181 181-181 179-181 179-181 169-181 177-177 173-177 179-181 181-185 179-181 181-181

Olieddu Oliva Bianca Oliva da polpa Oliva grossa Oliva Rossa Oliva torsa Oliva vera Olivago Olivastra di Populonia Olivastra Seggianese Olivastro d'Aprile Olivastro di Bucchianico Olivastro di Montenero Olivastro Dritto Olivastro Frentano Olivella appuntita Olivella di Cerchiara Olivella di Grottaminarda Olivetta Nera Olivo da olio Olivo da salare Olivo del mulino Olivo della Madonna Olivo delle Alpi Olivo di Casavecchia Olivo di Castiglione Olivo di San Lorenzo Olivoce Olivomensa Olivona Olivone di Viterbo OperaPia Orazio Orbetana Ornellaia Ortice Ortolana

214-222 218-222 214-222 214-214 208-218 212-214 214-214 212-212 212-222 212-218 222-222 208-222 218-222 212-212 208-222 214-222 212-218 214-218 212- 222 214-214 214-222 212-214 212-222 214-222 212-212 208-212 208-218 212-218 212-212 212-212 214-222 212-222 218-222 214-218 208-218 218-218 214-218

214-218 218-224 218-218 214-224 210-210 218-224 218-218 212-212 212-218 210-212 210-214 210-214 210-214 210-214 210-214 212-214 212-212 214-214 214-214 218-218 214-214 214-224 214-214 214-224 212-221 214-214 214-221 214-214 214-214 214-214 218-224 214-221 214-214 218-224 210-218 214-224 218-218

130-144 124-130 124-126 126-144 124-144 124-130 130-144 126-144 124-130 126-144 124-126 124-124 121-121 124-144 126-144 126-144 124-144 124-126 124-130 124-130 126-130 130-144 124-144 126-144 124-124 124-144 144-144 124-126 124-126 124-126 124-130 124-144 126-144 124-144 126-144 126-144 121-124

170-184 150-157 150-157 136-184 136-159 136-136 150-157 150-157 150-150 150-159 159-159 159-170 170-170 159-159 170-170 170-184 136-159 150-157 150-150 150-157 150-157 159-159 170-170 184-203 150-157 159-184 150-150 136-159 159-184 157-157 157-157 136-184 159-170 184-197 136-159 150-157 170-184

144-150 144-144 140-150 140-140 140-140 144-144 144-144 144-144 150-150 150-150 144-144 140-140 144-144 144-144 140-140 140-140 144-144 150-150 144-144 140-144 140-140 150-150 140-140 150-150 150-150 144-144 150-150 140-140 144-144 140-140 150-150 150-150 144-144 140-140 144-144 140-140 140-140

143-143 135-135 150-150 150-150 135-150 135-135 135-135 135-135 182-182 150-150 135-182 182-182 135-166 150-150 182-182 143-143 143-143 135-166 150-150 143-150 135-135 182-182 150-150 150-150 143-143 150-150 150-150 182-182 143-143 150-150 135-143 150-150 143-143 135-135 150-150 135-166 143-143

177-193 166-193 164-182 166-166 177-177 164-164 177-177 166-166 156-164 156-164 166-166 177-193 177-193 177-193 177-177 164-177 166-177 166-177 177-193 164-177 166-177 156-193 177-177 166-177 164-166 166-193 166-193 166-193 166-177 166-166 166-177 166-193 177-177 166-177 164-182 166-182 164-164

161-205 205-205 161-161 150-161 143-182 143-210 154-154 161-210 156-161 156-182 205-205 154-210 210-210 154-210 154-210 154-161 143-182 150-161 154-205 182-182 154-154 150-161 182-210 161-161 161-205 154-182 161-205 154-205 150-150 143-143 143-182 154-154 150-205 182-210 182-182 161-205 161-161

205-205 200-220 213-213 205-205 164-185 205-220 213-213 185-185 220-232 205-205 213-213 213-213 170-213 205-205 205-232 205-205 213-213 200-200 205-205 213-213 200-200 213-213 205-205 200-200 200-200 213-213 200-220 213-232 209-209 213-213 185-185 209-209 209-209 200-200 205-205 164-164 205-213

172-194 172-206 172-198 198-206 172-194 172-194 198-206 172-198 182-206 182-206 162-172 198-210 162-182 166-172 182-206 162-194 198-206 182-206 162-188 172-198 162-194 176-176 162-206 162-198 182-194 198-206 162-198 198-210 182-198 182-198 172-182 162-172 198-198 162-182 198-198 194-198 162-194

181-185 177-181 177-185 179-179 181-181 181-181 179-185 177-181 181-181 179-185 173-175 179-181 173-177 163-177 177-181 179-185 177-179 177-181 173-183 179-185 179-181 177-181 177-177 179-185 179-181 181-185 181-181 177-181 179-181 179-179 177-179 179-181 179-181 177-179 185-185 177-185 177-181

Ottobratica Ottobrina Paesana Bianca Paesana Nera Palma Palmarola Pampagliosa Paschixedda Pasola Passulunara Patrinostraru Pendolino 1 Pendolino 2 Pendolino 3 Pendolino 4 Pendolino 5 Pennulara Peppino Leo Peranzana Perciasacchi Pescarese Pesciatino Pezz'e quaddu Piangente 1 Piangente 2 Piangente 3 Piantone di Falerone Piantone di Macerata Piantone di Mogliano Pidicuddara Pignola Pirunara 1 Pirunara 2 Pisciottana Pizz'e Carroga Pizzo di corvo 1 Pizzo di corvo 2

212-222 214-214 208-212 212-222 214-222 208-222 214-214 214-222 208-218 212-222 212-212 212-222 212-222 222-222 212-222 212-222 212-222 208-218 212-222 212-214 208-212 208-208 214-214 208-212 212-222 208-212 218-222 212-222 214-214 212-212 214-222 212-222 212-222 214-222 212-222 212-212 212-212

212-212 218-224 214-224 214-214 218-221 214-224 218-224 212-221 214-214 210-210 210-214 214-214 214-214 214-214 214-214 214-214 214-214 214-214 212-214 214-224 214-224 212-214 214-221 214-214 214-214 214-224 218-218 218-224 218-224 210-210 214-214 214-214 214-214 218-218 218-218 210-214 210-214

124-124 126-144 124-144 124-130 130-144 124-144 124-130 126-144 124-144 124-144 130-144 126-130 126-130 126-130 126-130 126-130 130-144 124-144 130-144 124-144 124-126 126-144 126-144 126-126 124-130 144-144 124-130 124-144 124-130 124-144 126-130 124-126 124-126 121-124 126-144 124-144 124-144

157-170 159-170 159-184 150-150 170-184 150-157 170-184 157-159 150-157 159-184 157-170 150-159 150-150 150-150 150-150 150-150 150-159 150-159 170-170 136-136 159-170 170-184 136-203 157-184 150-150 159-184 150-157 170-184 159-159 150-184 150-157 150-170 150-170 136-150 170-184 136-159 136-159

140-150 150-150 144-144 144-144 144-150 144-144 140-140 144-150 144-144 144-144 144-144 144-144 144-144 144-144 144-144 144-144 140-150 144-144 144-144 150-150 144-144 144-144 144-150 144-144 144-144 144-144 144-144 144-144 144-144 144-144 150-150 144-144 144-144 140-140 140-140 144-144 140-140

143-143 150-150 150-150 150-182 135-135 143-143 143-143 143-143 143-143 150-150 150-150 166-166 150-166 166-166 150-166 166-166 143-143 135-135 182-182 143-143 143-143 143-182 135-135 150-150 150-150 150-150 135-135 143-143 135-135 150-150 143-143 150-150 150-150 143-143 135-135 150-150 150-150

177-193 166-193 177-177 177-193 177-193 166-166 164-182 164-177 177-177 166-193 166-193 177-177 177-177 177-182 177-182 177-177 177-193 177-182 166-193 164-177 177-193 164-166 166-166 166-193 177-193 166-193 166-177 166-193 166-166 166-166 177-193 177-177 177-177 177-182 164-193 166-166 166-166

143-154 150-205 182-210 154-205 161-205 182-210 143-143 150-161 150-182 182-182 143-182 154-205 154-205 154-205 154-205 154-205 143-182 143-205 143-205 161-182 154-205 150-182 150-205 154-154 154-205 182-210 210-210 182-210 182-210 182-182 154-182 182-182 182-182 150-161 150-150 143-182 143-182

170-205 200-220 205-205 213-213 205-205 200-200 200-200 213-213 170-170 108-205 205-232 205-205 205-213 205-205 205-205 205-213 209-220 108-108 108-205 213-220 220-243 220-232 213-213 205-205 205-205 205-205 205-232 185-200 200-232 205-205 205-205 205-205 205-205 213-232 125-125 213-213 213-213

182-206 182-194 162-166 162-204 172-172 194-198 182-206 176-182 198-206 176-182 182-198 162-206 162-206 162-206 162-206 162-206 198-206 172-194 162-172 162-198 198-206 162-194 162-162 162-206 162-206 162-206 182-182 162-198 198-206 162-182 182-206 162-182 162-182 172-182 162-172 162-198 172-182

177-185 181-181 163-177 177-177 181-185 177-179 181-181 179-181 177-177 179-179 173-179 181-181 181-181 181-181 181-181 181-181 179-179 177-177 181-181 181-181 179-185 179-185 181-185 181-181 181-181 181-181 177-185 179-181 181-181 177-179 181-185 179-181 179-181 177-181 179-185 169-185 175-179

Pizzutella Pocciolo Posola Posolella Precoce Procanica Provenzale Puntella Punteruolo Puntino Racemo 1 Racemo 2 Racioppa Racioppa Racioppella Raggiola Raja Raja Sabina Rajo Rastellina Ravece Raza Razzaio Razzo 1 Razzo 2 Razzola Reale Regina Resciola di Venafro Riminino Ritonnella Rizza Rizzitella Roma Romanella Romanella Romanella Molisana

208-222 218-222 212-218 214-218 208-212 214-222 218-218 222-222 212-214 208-212 212-212 208-208 208-208 212-222 218-222 212-222 214-222 212-222 214-218 214-222 214-218 212-222 212-218 208-212 208-212 214-222 212-222 208-212 208-222 214-222 214-218 212-222 214-218 218-222 208-222 212-212 212- 222

210-214 218-218 214-214 214-214 214-214 218-224 218-218 214-214 210-224 214-224 210-224 210-224 212-214 214-214 212-224 218-224 218-218 218-224 218-218 214-214 212-224 214-224 210-214 214-224 210-214 218-224 218-224 214-224 214-214 214-224 212-218 214-214 212-214 214-214 214-218 212-218 214-214

124-124 124-130 124-144 126-144 126-126 126-144 126-144 124-130 124-144 124-144 124-144 124-144 124-130 124-124 124-130 126-144 124-130 124-144 124-144 130-130 124-144 126-144 124-130 124-144 124-130 126-144 126-144 124-130 126-144 126-144 126-144 126-144 124-144 126-144 124-144 124-130 130-126

157-170 150-150 170-184 159-159 159-170 157-170 157-159 159-159 136-159 159-184 136-159 136-159 159-170 157-157 170-184 157-170 150-157 157-159 157-159 159-170 136-136 159-170 157-157 159-170 150-170 159-170 157-170 150-184 136-170 157-157 150-157 159-159 150-157 159-184 159-170 136-170 150-150

144-144 140-140 144-144 144-144 140-140 144-144 144-150 144-144 144-144 144-144 140-140 140-140 144-144 144-144 144-144 144-144 140-140 144-144 140-140 140-140 144-150 150-150 140-140 144-144 140-144 150-150 144-150 140-140 140-140 144-144 140-150 144-144 140-140 144-144 144-144 150-150 140-140

150-150 135-135 135-166 150-150 150-150 143-143 143-150 202-202 135-166 150-150 135-166 135-166 143-143 143-150 135-135 150-150 135-143 150-150 135-135 143-143 135-135 143-143 150-150 150-150 150-150 150-150 143-143 150-150 143-143 143-182 143-143 150-150 143-143 143-143 143-143 143-143 143-143

166-166 166-182 166-193 177-193 166-193 177-182 164-164 166-193 177-193 177-177 166-193 166-193 166-193 166-177 164-182 177-182 177-193 177-182 166-193 166-182 164-177 177-182 166-193 177-177 177-177 177-182 166-177 177-193 177-193 166-177 193-193 166-182 177-177 177-193 177-182 166-193 177-193

143-154 154-205 154-205 154-205 182-182 182-210 143-210 154-205 143-182 182-205 143-210 143-210 161-205 150-154 143-143 161-182 161-161 182-210 182-182 210-210 143-150 182-210 154-205 154-205 154-205 182-205 182-210 182-205 143-154 143-182 154-161 143-210 161-161 150-205 182-205 143-161 154-205

164-164 205-205 213-213 213-213 205-232 185-185 200-220 213-213 220-220 205-205 170-170 170-170 220-232 205-220 205-220 185-185 213-232 185-185 205-205 205-205 205-205 185-185 205-232 205-205 205-205 200-200 185-185 205-205 213-213 205-205 213-220 125-200 213-232 209-209 200-200 209-220 213-243

176-182 176-206 182-206 194-206 172-198 182-206 194-206 194-198 162-206 172-206 172-194 172-194 172-182 182-206 194-198 182-206 182-206 176-198 198-206 176-182 162-194 182-206 182-206 182-206 182-206 176-206 182-206 162-198 176-206 176-198 198-206 182-198 172-194 198-198 172-172 182-198 198-210

179-181 181-185 173-181 173-185 169-179 179-181 179-181 177-185 177-181 175-181 181-181 181-181 179-185 181-185 177-179 179-181 179-181 179-181 179-181 179-181 181-185 179-179 181-181 181-185 181-185 181-181 179-181 181-185 177-181 177-181 177-181 179-181 177-179 179-181 179-185 179-179 181-181

Rosciola Rosciola Coltodino Rosciola di Rotello Rosciola di Venafro Rosciola laziale Rosino Rossellino Rossello Rossina Rotondella Rotondella di Foggia Rotondella di Sanza 1 Rotondella di Sanza 2 Rumignana Rustica Ruveia Salegna di Larino Salella Salicino Salvia Salviana Sammartinara Sammartinenga San Benedetto San Felice Acquasparta 1 San Felice Acquasparta 2 San Felice Acquasparta 3 San Felice Acquasparta 4 San Felice Acquasparta 5 San Francesco San Lazzero Sanginara Santa Caterina Santa Maria Sant'Agatese Sant'Agostino Santomauro

214-222 208-218 208-212 222-222 212-222 222-222 208-208 208-212 214-222 212-222 208-218 212-218 212-218 208-212 212-222 214-214 212-222 214-218 212-222 218-222 214-222 212-212 208-222 208-212 212-212 208-212 212-222 212-212 212-212 208-208 208-212 214-222 208-208 214-218 208-222 212-212 208-222

218-218 212-212 214-224 214-214 218-224 212-214 210-212 214-214 214-214 214-214 214-214 218-218 218-218 214-214 210-214 214-218 214-214 214-214 210-214 218-224 214-224 214-214 214-214 214-214 212-212 214-214 214-224 214-259 214-259 210-210 214-224 218-218 210-212 214-218 214-214 210-214 214-214

126-144 126-144 124-144 124-124 144-144 124-130 126-144 124-130 126-144 130-144 124-126 126-144 126-144 121-121 124-126 124-126 124-130 124-130 124-126 126-144 130-144 126-144 124-144 124-144 130-144 126-144 124-144 130-144 130-144 126-130 124-144 124-126 124-144 124-144 124-126 124-144 124-144

184-197 184-184 159-184 150-184 157-157 184-184 170-184 184-184 150-157 150-150 150-157 157-159 157-159 150-170 150-184 170-184 150-150 136-150 136-159 150-159 150-157 159-159 136-159 157-157 170-170 157-157 184-184 159-184 159-184 170-184 159-184 150-157 136-197 157-159 150-184 170-170 157-184

144-150 150-150 144-144 144-144 144-150 140-140 144-144 144-144 150-150 140-140 140-140 140-140 140-140 140-140 140-140 140-140 140-140 140-140 144-144 144-150 140-140 144-144 144-144 144-144 144-144 144-144 144-144 140-140 140-140 140-140 144-144 140-140 150-150 140-150 144-144 144-144 144-150

143-143 182-202 150-150 202-202 143-143 150-150 202-202 150-150 135-135 150-150 135-135 143-150 143-150 150-150 135-135 143-143 150-150 143-143 150-150 143-143 135-135 150-150 143-143 135-143 150-150 150-150 150-150 150-150 150-150 150-150 150-150 143-143 143-143 143-143 150-150 143-143 143-143

177-182 177-182 177-177 177-193 177-193 156-193 164-166 177-177 177-193 166-166 177-177 164-182 164-182 166-193 166-166 164-182 177-193 164-177 156-193 177-193 177-182 177-177 166-177 166-193 166-193 166-193 166-193 166-193 166-193 156-182 177-177 164-177 156-164 164-177 177-177 166-166 166-182

154-205 156-161 182-182 154-205 182-210 154-205 156-205 154-182 161-210 154-182 154-182 150-205 150-205 182-182 154-205 154-205 154-205 150-182 182-205 154-210 182-210 182-182 143-182 154-182 154-205 143-154 182-205 182-182 182-182 182-182 182-205 154-161 150-182 154-205 182-182 154-154 143-205

205-205 200-200 205-205 243-243 185-220 220-243 220-232 213-213 213-232 200-200 185-205 205-232 205-232 220-220 205-205 213-232 220-243 213-232 205-205 185-185 185-220 205-205 213-213 205-205 205-205 205-232 205-205 220-220 220-220 213-213 205-205 205-205 220-220 205-205 213-213 108-108 205-232

198-206 182-206 162-166 198-206 162-198 182-206 172-194 176-194 176-182 182-198 182-198 198-206 198-206 172-206 198-206 172-194 198-210 172-182 162-198 172-182 182-206 162-162 162-176 162-198 198-206 198-206 198-206 198-206 198-206 182-206 172-194 182-206 162-194 172-172 172-182 172-198 182-198

177-179 179-181 163-177 177-181 179-181 185-185 179-185 177-185 177-181 179-181 177-179 179-181 179-181 169-179 173-173 181-185 173-181 177-181 179-181 179-181 181-185 179-181 179-181 179-185 177-179 177-179 177-179 177-179 177-179 179-181 179-181 181-185 185-185 173-179 177-179 177-181 177-179

Sargano Sargano Scarpetta Selvatico Semidana Sessana Simona Sinopolese Sirole Sivigliana da olio Spagnola di Missano Sperone di Gallo Spezzanese Spinoso Taggiasca Tenacella Tendellone Termite di Bitetto Terza grande Terza piccolo Toccolana Tombarello Tonda di Alife Tonda di Cagliari Tonda di Filadelfia Tonda di Filogaso Tonda di Strongoli Tonda Dolce Tonda Dolce di Partanna Tonda iblea Tondina Tortiglione Toscanina Tunnulidda Turdunazza antimosca 1 Turdunazza antimosca 2 Uccellara

214-222 218-222 208-218 214-222 214-214 214-214 212-222 212-214 214-222 214-222 208-208 212-212 212-222 208-222 208-212 218-218 214-222 208-218 214-222 214-222 212-222 212-212 214-214 212-212 212-222 212-212 212-222 208-208 212-212 208-212 208-222 208-212 218-218 208-212 212-212 212-212 212-212

218-224 214-214 212-221 214-214 214-218 214-224 210-210 212-212 218-224 214-214 212-224 214-214 214-214 214-214 214-224 214-218 218-221 214-214 214-221 212-218 214-214 212-212 214-214 214-221 212-218 212-250 212-218 214-214 214-214 210-214 214-214 214-224 210-214 214-214 210-214 210-214 212-212

124-144 130-144 124-124 126-144 130-144 126-130 124-126 124-130 130-144 130-130 124-144 130-130 124-130 126-144 124-144 126-144 126-144 124-144 126-144 126-144 124-130 130-144 124-126 126-144 126-144 124-124 124-126 126-144 144-144 124-144 126-144 124-126 124-144 126-144 126-144 126-144 124-144

159-184 150-157 150-159 150-157 136-184 170-184 159-159 170-184 150-157 159-159 157-159 170-170 136-170 159-184 159-170 157-159 159-159 150-159 157-159 157-159 150-150 150-159 150-157 170-203 150-157 157-170 136-150 150-159 157-184 136-159 159-159 159-170 150-159 159-184 159-184 159-184 159-170

144-144 150-150 144-144 150-150 144-150 144-144 144-144 144-144 140-140 150-150 144-144 140-140 144-144 144-144 144-144 140-150 140-140 144-144 144-150 144-150 144-144 150-150 150-150 150-150 144-144 150-150 144-144 144-144 140-140 144-144 150-150 144-144 144-144 144-144 140-140 140-140 144-144

143-143 135-135 150-150 135-135 135-135 143-143 166-166 143-143 135-135 135-135 143-143 143-143 143-143 150-150 143-143 143-143 150-150 143-143 143-143 143-143 150-182 143-143 166-166 135-135 143-143 143-143 143-143 143-143 150-150 150-150 143-143 166-166 143-143 150-150 150-150 150-150 143-143

177-182 166-177 177-182 177-193 164-193 166-177 166-177 177-193 177-182 166-177 166-177 166-193 166-193 166-182 177-182 164-164 166-182 177-182 164-177 164-177 177-193 177-193 164-193 166-193 166-193 166-166 166-193 166-193 166-193 166-193 166-182 177-177 177-177 166-193 166-193 166-193 166-177

182-210 154-205 143-143 161-210 150-161 182-205 154-154 154-210 182-210 210-210 143-210 182-182 143-154 143-210 182-205 161-161 143-182 143-205 150-161 150-161 154-205 143-182 150-154 143-143 143-205 143-182 143-182 143-154 182-182 154-154 143-182 154-210 161-182 182-182 143-143 143-143 143-143

185-185 200-205 200-200 213-232 200-200 213-213 108-170 164-205 185-220 213-232 185-185 213-213 205-205 205-213 205-205 213-213 205-205 108-170 213-213 213-213 205-205 205-205 205-205 213-220 205-232 205-205 205-220 170-170 205-205 108-205 205-232 220-243 164-164 205-205 213-232 213-232 200-200

182-198 172-182 172-198 172-182 162-172 182-206 182-182 162-198 176-206 162-176 176-198 162-172 162-198 194-198 172-194 172-206 176-198 172-194 182-198 176-182 162-188 172-206 182-206 162-172 162-198 198-206 162-198 162-198 182-182 162-198 198-206 198-206 172-198 172-182 172-172 172-198 198-206

179-181 177-177 179-181 177-181 181-185 181-185 175-181 177-179 181-185 181-181 181-185 169-179 179-185 179-181 179-181 181-181 179-181 179-179 179-181 179-181 173-183 177-177 181-185 185-185 177-185 177-179 177-185 177-185 177-181 169-177 177-179 179-181 177-181 169-181 173-179 179-185 179-181

Ugea di Savoia Ugliastride Vaddara Vallanella 1 Vallanella 2 Verdello Vigna della Corte Vocio Vraja Zaituna Zimbimbo Zinzifarica

212-212 208-218 208-212 208-212 208-212 212-222 212-214 214-214 208-218 212-212 208-218 212-222

212-212 214-221 210-214 214-224 210-224 210-214 214-218 214-214 214-214 214-214 214-224 214-214

126-130 124-144 124-124 130-144 124-144 124-126 124-144 130-144 126-144 124-144 124-144 126-144

159-184 170-184 157-159 170-170 157-157 159-170 157-159 150-157 170-184 150-170 157-170 150-159

140-140 140-150 144-144 144-144 144-144 144-144 140-140 140-140 144-144 140-140 144-144 144-144

143-143 143-143 150-150 150-150 150-150 150-150 143-150 143-143 143-143 150-150 135-135 143-143

166-166 166-193 177-193 166-193 166-193 166-166 164-177 166-177 166-193 166-193 177-182 177-177

143-161 143-182 154-182 143-210 154-154 143-143 150-154 161-205 143-182 182-182 182-182 143-182

205-205 200-200 164-205 213-232 205-205 170-220 213-220 205-213 200-200 108-108 108-108 205-205

162-172 172-176 198-198 162-206 162-206 182-206 182-206 198-206 162-198 182-206 198-198 162-198

177-181 185-185 179-179 173-175 173-175 181-181 179-181 177-181 179-185 177-177 177-179 177-179